Biological nitrogen fixation in grass–clover affected by animal excreta

Aiming at estimating the average N2-fixation in a pasture, ap preciating the great variability due to patchy urine and dung deposition, the in fluence of dairy cow excreta on biological N2-fixation in a perennial ryegrass–white clover mixture was studied using natural urine and dung. Application of urine as well as dung affected the N2-fixation by promoting the growth of grass and thereby the proportion of clover was significantly reduced. Also the proportion of clover-N derived from the atmosphere (pNdfa) was significantly reduced. In control plots clover dry matter constituted between 40 and 50% of the total dry matter production and the pNdfa ranged between 0.8 and 0.9. Addition of urine caused a significant increase in the grass growth rates, which was the primary reason for a decrease in proportion of clover. At the same time pNdfa decreased to 0.2–0.4 followed by an increase resulting in a total reduction of 45% in the N2-fixation in urine affected areas over a period of four months. The dung only affected the N2-fixation for a distance of up to 10 cm from the edge of the dung pats. In this border area the pNdfa decreased from 0.85 to 0.75 during one month after application followed by an increase, so that after three months there was no difference between pNdfa at 0–10 and 10–20 cm distance from the dung hill. The proportion of clover was lower in the 0–10 cm than in the 10–20 cm distance, which totally resulted in a total reduction of 20% in the N2-fixation over a period of four months in the 0–10 cm area around the dung pats. Considering the proportion of a pasture which may by affected by excreta at a stocking density of 4–6 cows ha-1, the length of the grazing period, the frequency of excretion and the area covered by individual patches, it was estimated that the N2-fixation in a grass-clover pasture would be reduced by 10–15% compared to the N2-fixation in a grass-clover sward not exposed to animal excreta.     

A field study of nitrogen dynamics and spring barley growth as affected by the quality of incorporated residues from white clover and ryegrass

Biofumigation refers to the suppression of soil-borne pests and pathogens by biocidal compounds released by Brassicaceous green manure and rotation crops when glucosinolates (GSLs) in their tissues are hydrolysed. We investigated the effect of environment and ontogeny on the GSL production, and thus biofumigation potential, of eight entries from five Brassica species. The environments included autumn and spring sown field plots (FA and FS) and potted plants grown under ambient conditions (PAM) or in a temperature controlled glasshouse at 20 °C/12 °C (PTC). GSL concentration was measured in the root and shoot tissue at buds-raised, flowering and maturity. Of particular interest was the suitability of the pot-grown plants for screening large numbers of brassicas for GSL production. The type of GSLs present in the tissues and their relative proportions remained relatively constant across environments and at different growth stages, with the exception of an increase in indolyl GSLs in the FS environment suspected of being induced by insect attack. Total GSL concentration generally declined from buds-raised to flowering in all environments, and was lowest at maturity. The exceptions were B. campestris, which had higher GSL concentration at flowering than at buds-raised, and the PTC environment in which most species also showed an increase at flowering. Despite GSL types and their proportions remaining relatively constant, the total GSL concentration in the root and shoot tissue of all entries varied significantly with environment (3–10-fold) and was generally ranked FS>PAM>FA>PTC. Interactions between species and environments meant that the ranking of the Brassica entries for total shoot and root GSL concentration changed with environment. However within three entries from B. napus, the ranking was consistent across the environments. The added effect of environment on phenological development and biomass production further influenced GSL production (the product of GSL concentration and biomass) on a ground area basis. The results suggest that glasshouse environments can be used to determine the types and proportions of GSLs present, and to rank entries within, but not between species for the total concentration in the tissues. However the influence of the environment on both GSL concentration and biomass production suggests that an accurate estimate of GSL production on a ground area basis to assess biofumigation potential will require measurement in the target environment.     

Distribution of photosynthate produced before and after anthesis in tall and semi-dwarf winter wheat, as affected by nitrogen fertiliser

In 2 years the distribution of radioactivity recovered in entire shoots of field-grown winter wheat was determined at various times after exposing the top two leaves (flag leaf or second leaf) to ¹⁴CO₂ for 30 s. In 1976 when ¹⁴C was supplied to either leaf 14 days before anthesis, 30% was in the ear at anthesis. Less than 5% was in the leaf exposed to ^I4CO₂. The remainder was equally divided between the stem above and below the flag-leaf node when the flag leaf had been exposed, and was mainly in the lower part of the stem when the second leaf had been exposed. By maturity the proportion in the stem had decreased; 20% of the total activity was in the grain and 30% was still in the ear structures. When ¹⁴C was supplied 10 days after anthesis, the proportion in the ear 24 h later ranged from 42 to 69% of that in the whole shoot when the flag leaf was exposed, and from 6 to 28% when the second leaf was exposed. At maturity these proportions increased to 92 and 85% when the ¹⁴C had been supplied to flag leaves and second leaves respectively. When ¹⁴C was supplied 25 days after anthesis to either flag leaves or second leaves, more than 90% of the activity was in the mature ears. Less than 5% of the ¹⁴C remaining at maturity from any treatment was still in the leaf exposed to ¹⁴CO₂. Between 2 and 6% of ¹⁴C supplied after anthesis was in the non-grain parts of the ear. The proportion of the ¹⁴C in the ear was greatest for the semi-dwarf varieties Maris Fundin and Hobbit, less for Maris Huntsman, and least for Cappelle-Desprez. These varietal differences were large 24 h after exposure to ¹⁴CO₂, especially in 1976. They were small and rarely significant at maturity. Nitrogen fertiliser up to 210 kg N ha^-1 had negligible effects on the distribution of ¹⁴C, although it greatly increased growth and yield, especially in 1975.     

The influence of phosphorus and nitrogen on millet and clover growing in soils affected by salinity

Summary The growth of foxtail millet and clover in soils of varying degrees of salinity (0.5 to 13 mmhos/cm), treated with nitrogen and phosphorus, was studied. Salinity levels were achieved by addition of sodium chloride. Nitrogen (10 to 60 ppm N) and phosphorus (6.4 to 44.8 ppm P) were added as NH₄NO₃ and H₃PO₄, respectively. The growth of millet decreased sharply with increase in soil salinity, when N-P treatments were not applied. The development of this plant altered under saline conditions, however, when nitrogen and phosphorus were added; various N-P combinations affected plant growth in saline soil differently. Phosphorus, when applied at relatively high rates, significantly improved plant growth. Increased rates of nitrogen in the N-P treatments generally had no significant effect on growth; it was reduced when the N/P ration was highest. Clover ceased to grow when the salinity of the soil exceeded 7 mmhos/cm and no N-P was added. Phosphorus enhanced the growth of clover, and at high rates of its application in the N-P combinations, clover grew even at the highest salinity level tested (13 mmhos/cm). Nitrogen increase had no marked effect on plant development. Comparing enhancement of growth at high salinity levels, clover was somewhat less affected by the N-P treatment than millet. The top/root ratio of clover generally increased with increase of phosphorus in the N-P combinations. This research (Parts I and II) was supported in part by a grant from the U.S. Department of Agriculture under P. L. 480.     

Competition for nutrients between grasses and white clover

Summary Competition between three grass species and white clover for phosphorus and potassium were studied in pot trials. Differences in the ability of the individual grasses to compete with clover were noted to be of the same order for both nutrients.Nitrogen application was shown to increase competition for nutrients by grasses in clover associations without postulating an indirect mechanism acting through competition for light.Evidence of a light by nutrient competition interaction was observed and its mechanism discussed.     

Effect of white clover cultivar on apparent transfer of nitrogen from clover to grass and estimation of relative turnover rates of nitrogen in roots

The apparent transfer of N from clover to associated grass was evaluated over a four year period both on the basis of harvested herbage and by taking account of changes in N in stubble and root (to 10 cm depth) in swards with perennial ryegrass and three different white clover cultivars differing in leaf size. The large leaved Aran transferred 15% of its nitrogen while Huia transferred 24% and the small leaved Kent Wild White transferred 34%. When changes in stubble and root N were taken into account the percentage of N transferred was calculated to be 5% less than in harvested herbage only, as the small leaved types had proportionately more N in the roots and stolons, but the large leaved type was probably more competitive towards the grass.Loss of N from clover roots from July to October was compared to that from grass roots in a grass/white clover sward continuously stocked with steers using a method which incorporated tissue turnover and ¹⁵N dilution techniques. Less than 1 mg N m^-2 d^-1 was lost from the grass roots. In contrast 8 mg m^-2 d^-1 were estimated to be lost from clover roots while 12 mg N m^-2 d^-1 were assimilated.It is concluded that clover cultivar and competitive ability on grass have to be taken into account together with the relationship between N turnover in roots and N available for grass growth when modelling N transfer in grass/clover associations.     

Residual nitrogen pools in mature winter wheat straw as affected by nitrogen application

Plant and Soil - Reduction of the amount of nitrogen (N) remaining in mature wheat straw is an essential challenge in order to boost grain protein and N use efficiency. However, very limited...     

Stepwise incorporation of white clover (Trifolium repens L.) as fertiliser increases nitrogen fixation and improves nitrogen retention when intercropped with leek (Allium porrum L.)

Plant and Soil - We investigated whether the incorporation of intercropped white clover as a split-dose fertiliser improves N retention of the plant-soil system in leek production. White clover and...     

The absorption and transport of manganese by perennial ryegrass and white clover as affected by silicon

Summary The effects of added silicon on the absorption and transport of manganese in perennial ryegrass (an accumulator of Si) were determined and compared with those found in white clover (which has restricted Si uptake). The plants were grown in flowing solution culture in two experiments with Si in the nutrient solution maintained at 0, 10 or 20 mgl⁻¹ and Mn at 0.005 mgl⁻¹. By the final harvests, the plants contained concentrations of both Mn and Si that were comparable to those found in field-grown plants. In common with other findings, white clover had very much lower concentrations of Si in both shoots and roots than did ryegrass but there was no effect of Si treatment on the growth of either species. In both species, the concentrations of Mn were initially greater in roots than in shoots, but values in both plant parts decreased with time and by the final harvests, were similar. The rates of absorption of Mn by roots and its subsequent transport to shoots were also similar in both species. In contrast to findings for other species in other studies based on conventional solution culture, there was no effect of added Si on either absorption or transport of Mn in clover or ryegrass. It was therefore concluded that any effect of Si on the behaviour of Mn in plants must result from changes in distribution and partitioning within leaf tissues and cells.     

Mycoplasma development and cell alterations in white clover affected by clover dwarf an electron microscopy study

Summary The presence of mycoplasma has been demonstrated in the phloem of leaves of white clover (Trifolium repens L.) affected by clover dwarf. Mycoplasma-like bodies were found both in parenchyma and companion cells and in sieve elements.In young parenchyma and companion cells mycoplasma-like bodies appeared as round or oval particles with high ribosomal content, delimited by a ribosome-bearing membrane. Their diameter ranged between 50 and 400 nm. In mature sieve elements they were larger, more pleomorphic, and showed a central clear area containing presumed DNA filaments. Budding and dividing forms were sometimes seen among them.The main alterations found in the infected cells were: increased ribosome content, dilation of the perinuclear space, degeneration of mitochondria and chloroplasts, and cytoplasmic vacuolation. Many cells appeared completely disrupted and their content was replaced by a great number of pleomorphic mycoplasma.This investigation was supported by a grant of Consiglio Nazionale delle Ricerche, Rome.     

Growth and solute composition of the salt-tolerant kallar grass [Leptochloa fusca (L.) Kunth] as affected by nitrogen source

A study was conducted to elucidate the effect of N form, either NH₄ ⁺ or NO₃ ^–, on growth and solute composition of the salt-tolerant kallar grass [Leptochloa fusca (L.) Kunth] grown under 10 mM or 100 mM NaCl in hydroponics. Shoot biomass was not affected by N form, whereas NH₄ ⁺ compared to NO₃ ^– nutrition caused an almost 4-fold reduction in the root biomass at both salinity levels. Under NH₄ ⁺ nutrition, salinity had no effect on the biomass yield, whereas under NO₃ ^– nutrition, increasing salinity from 10 mM to 100 mM caused 23% and 36% reduction in the root and shoot biomass, respectively. The reduced root growth under NH₄ ⁺ nutrition was not attributable to impaired shoot to root C allocation since N form did not affect the overall root sugar concentration and the starch concentration was even higher under NH₄ ⁺ compared to NO₃ ^– nutrition. The low NH₄ ⁺ (2 mM) and generally higher amino-N concentrations in NH₄ ⁺- compared to NO₃ ^–-fed plants indicated that the grass was able to effectively detoxify NH₄ ⁺. Salinity had no effect on Ca²⁺ and Mg²⁺ levels, whereas their concentration in shoots was lower under NH₄ ⁺ compared to NO₃ ^– nutrition (over 66% reduction in Ca²⁺; over 20% reduction in Mg²⁺), but without showing deficiency symptoms. Ammonium compared to NO₃ ^– nutrition did not inhibit K⁺ uptake, and the K⁺-Na⁺ selectivity either remained unaffected or it was higher under NH₄ ⁺ than under NO₃ ^– nutrition. Results suggested that while NH₄ ⁺ versus NO₃ ^– nutrition substantially reduced root growth, and also strongly modified anion concentrations and to a minor extent concentrations of divalent cations in shoots, it did not influence salt tolerance of kallar grass.     

Crop response to magnesium fertilization as affected by nitrogen supply

Background

Crop yield depends in large part on the availability and accessibility of nitrogen in the soil. For optimal yield, the soil nitrogen must be available at critical periods of crop development, and in a form that is accessible for plant uptake and use. Ancillary crop nutrients can alter the plant’s ability to access and utilize nitrogen. Therefore, crop fertilization with magnesium should focus on its effect on nitrogen management. This conceptual review aims to assess the present state of knowledge regarding the importance of magnesium in fulfilling both objectives.

Scope

The response to fertilizer magnesium of high-yielding wheat, maize, sugar beet and potato crops was evaluated using published and unpublished data on yield, yield components and nitrogen uptake. A simple, stepwise regression and path analysis was applied to explain the effect of fertilizer magnesium on yield and yield components.

Conclusions

The effect of soil or foliar applied magnesium on yield of crops was inconsistent due to (i) weather experienced during the growth season, (ii) rates of applied fertilizer nitrogen, and (iii) the (natural background levels of?) magnesium available in the soil. The yield increase due to magnesium application was related to the extra supply of nitrogen. In cereals, magnesium application resulted in a higher number of ears and/or thousand grain weight (TGW), stressing the magnesium-sensitive stages of yield formation. The increase of sugar beet yield was most pronounced in dry years. The main conclusion gleaned from the review underlines a positive effect of magnesium on nitrogen uptake efficiency. The optimal yield forming effect of fertilizer magnesium can generally occur under conditions of relatively low nitrogen supply (soil + fertilizer nitrogen), but high supply of magnesium. This phenomenon can best be described as “magnesium-induced nitrogen uptake”.     

Senescence and decomposition of white clover stolons in grazed upland grass/clover swards

The fate of marked sections of stolons of white clover (Trifolium repens) over a 50-week period from May 1987 was followed in grazed grass/clover swards maintained at 5-cm sward surface height with and without N fertiliser. There was little effect of N treatment on the pattern of survival of stolon sections. The proportion of live stolons recovered decreased during the experiment, and in May 1988 on average only 29% of the marked sections remained alive. At all harvests only a small percentage of stolon sections showed signs of senescence; the maximum percentage, on average 20% of those marked, occurred in autumn, 15–20 weeks after marking. Following this period, i.e. in late autumn/winter, the most rapid increase in percentage of decomposed stolons was measured. Over 50% of stolon sections were buried within the 5-week period following marking and nearly all were buried after 20 weeks; generally a much smaller proportion of stolon tips was buried. Nutrient concentrations of N, P and K fell to their lowest levels in autumn, before increasing in the following spring. Results are discussed in relation to the cycling of nutrients via stolon senescence.     

Use of white clover as an alternative to nitrogen fertiliser for dairy pastures in nitrate vulnerable zones in the UK: productivity, environmental impact and economic considerations

Perennial ryegrass and perennial ryegrass/white clover permanent dairy pastures are compared with respect to productivity, environmental impact and financial costs in nitrate vulnerable zones (NVZ) in the UK. With appropriate management, and utilisation of recommended perennial ryegrass and white clover cultivars, white clover is likely to stabilise at around 20% of total dry matter production in a mixed pasture. Plant dry matter production and milk production from a perennial ryegrass/white clover pasture are likely to be similar to that from a perennial ryegrass pasture receiving 200 kg N ha⁻¹ annum⁻¹ and around 70% of that obtained with perennial ryegrass supplied with 350–400 kg N ha⁻¹ annum⁻¹. Nitrate, phosphorus and methane losses from the system and decreases in biodiversity relative to a grazed indigenous sward are likely to be similar for a perennial ryegrass/white clover pasture and a perennial ryegrass pasture receiving 200 kg N ha⁻¹ annum⁻¹: nitrate leachate from both systems is likely to comply with European legislation. Greenhouse gas emissions resulting from nitrogen (N) fertiliser production would be avoided with the perennial ryegrass/white clover pasture. Within NVZ stocking rate restrictions, white clover can provide the N required by a pasture at a lower financial cost than that incurred by the application of N fertiliser.     

Elevated CO2 effects on carbon and nitrogen cycling in grass/clover turves of a Psammaquent soil

Effects of elevated CO₂ (525 and 700 L L^–1), and a control (350 L L^–1 CO₂), on biochemical properties of a Mollic Psammaquent soil in a well-established pasture of C3 and C4 grasses and clover were investigated with continuously moist turves in growth chambers over four consecutive seasonal temperature regimes from spring to winter inclusive. After a further spring period, half of the turves under 350 and 700 L L^–1 were subjected to summer drying and were then re-wetted before a further autumn period; the remaining turves were kept continuously moist throughout these additional three consecutive seasons. The continuously moist turves were then pulse-labelled with ¹⁴C-CO₂ to follow C pathways in the plant/soil system during 35 days.Growth rates of herbage during the first four seasons averaged 4.6 g m^–2 day^–1 under 700 L L^–1 CO₂ and were about 10% higher than under the other two treatments. Below-ground net productivity at the end of these seasons averaged 465, 800 and 824 g m^–2 in the control, 525 and 700 L L^–1 treatments, respectively.in continuously moist soil, elevated CO₂ had no overall effects on total, extractable or microbial C and N, or invertase activity, but resulted in increased CO₂-C production from soil, and from added herbage during the initial stages of decomposition over 21 days; rates of root decomposition were unaffected. CO₂ produced h^–1 mg^–1 microbial C was about 10% higher in the 700 L L^–1 CO₂ treatment than in the other two treatments. Elevated CO₂ had no clearly defined effects on N availability, or on the net N mineralization of added herbage.In the labelling experiment, relatively more ¹⁴C in the plant/soil system occurred below ground under elevated CO₂, with enhanced turnover of ¹⁴C also being suggested.Drying increased levels of extractable C and organic-N, but decreased mineral-N concentrations; it had no effect on microbial C, but resulted in lowered microbial N in the control only. In soil that had been previously summer-dried, CO₂ production was again higher, but net N mineralization was lower, under elevated CO₂ than in the control after autumn pasture growth.Over the trial period of 422 days, elevated CO₂ generally appears to have had a greater effect on soil C turnover than on soil C pools in this pasture ecosystem.     

Bacteriophage in relation to nitrogen fixation by red clover

Summary Cultures of Rhizobium trifolii resistant to the action of a given bacteriophage, may vary appreciably in their nitrogen fixing ability in association with the proper host plant. Likewise, cultures of Rh. trifolii, sensitive to bacteriophage may or may not benefit the host plant as judged by nitrogen fixation.Since sensitive and resistant cultures may be comparable in their nitrogen fixing capacity, it appears that the behavior of a culture of Rh. trifolii toward the lytic action of a specific bacteriophage in vitro cannot be correlated with its nitrogen fixing ability in association with the host plant.If a bacteriophage is added to a sensitive strain of Rh. trifolii used for inoculation of red clover, the cultures recovered from the nudules are often only of the resistant type. When this occurs the nitrogen fixed through association of plant and bacteria is decreased in the case of a good strain, but is unaffected if the strain is of the poor type. The addition of phage to a resistant strain of Rh. trifolii used for inoculation of red clover plants does not change the resistant type of culture recovered from the nodules, nor is the fixation of nitrogen by plant and bacteria affected.     

Acetylene reduction assay on white clover genotypes and on grass/clover swards

Acetylene reduction assay was used to measure the nitrogenase activity of white clover genotypes in pots and of grass/clover swards in situ. Much of the variation in nitrogenase activity of single genotypes and hybrid populations was associated with plant dry weight. After adjustment for plant dry weight it was concluded that there was limited scope for selection for increased nitrogenase activity. In plant breeding this technique would seem to have greatest application in the selection for continued nitrogen fixation activity in the presence of inhibitory factors such as high levels of mineral nitrogen. The in situ studies revealed differences in nitrogenase activity of grass/clover swards based on contrasting cultivars of white clover. These differences were due to variation in clover density and also to variation in activity per unit clover dry weight. It was concluded that the variation in nitrogenase activity per unit dry weight reflected differences in growth pattern of the cultivars in the autumn when these assays were made. The in situ studies offer a means whereby the nitrogenase activity of cultivars and selected families can be monitored under varying levels of mineral nitrogen and other husbandry treatments without disruption of the sward.     

Phosphorus accumulation in grains of japonica rice as affected by nitrogen fertilizer

Background and aims

This study aims to investigate the effect of nitrogen (N) on grain phosphorus (P) accumulation in japonica rice.

Methods

Six cultivars with contrasting agronomic traits were grown for 3 years (from 2008 to 2010) of field experiments under seven N treatments and 1 year (in 2010) of pot experiments with five N treatments to study the effect of N on grain phosphorus accumulation and to explore its physiological foundation.

Results

Grain total P and phytic acid concentration showed a clearly decreasing trend as N rate increased for both field and pot experiments. Pot experiment revealed that application of N increase plant biomass, but tended to lower plant P uptake, especially for the split topdressing treatments. Both harvest index (HI) and P harvest index (PHI) increased with N rate, but PHI was consistently higher than HI, indicating the larger proportion of P translocation to grain than that of dry matter by N. Further, ratio of PHI/HI differed significantly among genotypes, but was stable across contrasting N treatments.

Conclusions

The combination of decreased plant P uptake and dilution effect of increased grain yield by N is proposed as underlying mechanism of the decreased grain P concentration by high N.     

Short-term immobilisation and crop uptake of fertiliser nitrogen applied to winter wheat: effect of date of application in spring

Previous studies on the fate of fertiliser nitrogen applied to winter wheat in temperate climates have shown that nitrogen (N) applied early, at tillering for wheat, was less efficiently taken up than N applied later in the growth cycle. We examined the extent to which the soil microbial N immobilisation varied during the wheat spring growth cycle and how microbial immobilisation and plant uptake competed for nitrogen. We set up a pulse-15N labelled field experiment in which N was applied at eight development stages from tillering (beginning of March) to anthesis (mid-June). Each application was 50 kg N ha-1 as 15N labelled urea except for the first application which was 25 kg N ha-1. The distribution of fertiliser 15N in shoots, roots, mineral and organic soil N was examined by destructive sampling 7 and 14 days after each 15N pulse. The inorganic 15N pool was almost depleted by day 14. The N uptake efficiency increased with later applications from 45% at tillering to 65% at flowering. N immobilisation was rather constant at 13–16% of N applied, whatever the date of application. The increase in plant 15N uptake resulted in an increase in the total 15N recovery in the plant-soil system (15N in soil +15N in plant), suggesting that gaseous losses were lower at the later application dates.