Intercropping with pulses to concentrate nitrogen and sulphur in wheat

The effects of intercropping wheat with faba bean (Denmark, Germany, Italy and UK) and wheat with pea (France), in additive and replacement designs on grain nitrogen and sulphur concentrations were studied in field experiments in the 2002/03, 2003/04 and 2004/05 growing seasons. Intercropping wheat with grain legumes regularly increased the nitrogen concentration of the cereal grain, irrespective of design or location. Sulphur concentration of the cereal was also increased by intercropping, but less regularly and to a lesser extent compared with effects on nitrogen concentration. Nitrogen concentration (g/kg) in wheat additively intercropped with faba bean was increased by 8% across all sites (weighted for inverse of variance), but sulphur concentration was only increased by 4%, so N:S ratio was also increased by 4%. Intercropping wheat with grain legumes increased sodium dodecyl sulphate (SDS)-sedimentation volume. The effect of intercropping on wheat nitrogen concentration was greatest when intercropping had the most deleterious effect on wheat yield and the least deleterious effect on pulse yield. Over all sites and seasons, and irrespective of whether the design was additive or replacement, increases in crude protein concentration in the wheat of 10 g/kg by intercropping with faba bean were associated with 25-30% yield reduction of the wheat, compared with sole-cropped wheat. It was concluded that the increase in protein concentration of wheat grain in intercrops could be of economic benefit when selling wheat for breadmaking, but only if the bean crop was also marketed effectively.

Pea-barley intercropping and short-term subsequent crop effects across European organic cropping conditions

Grain legumes are known to increase the soil mineral nitrogen (N) content, reduce the infection pressure of soil borne pathogens, and hence enhance subsequent cereals yields. Replicated field experiments were performed throughout W. Europe (Denmark, United Kingdom, France, Germany and Italy) to asses the effect of intercropping pea and barley on the N supply to subsequent wheat in organic cropping systems. Pea and barley were grown either as sole crops at the recommended plant density (P100 and B100, respectively) or in replacement (P50B50) or additive (P100B50) intercropping designs. In the replacement design the total relative plant density is kept constant, while the additive design uses the optimal sole crop density for pea supplementing with 'extra' barley plants. The pea and barley crops were followed by winter wheat with and without N application. Additional experiments in Denmark and the United Kingdom included subsequent spring wheat with grass-clover as catch crops. The experiment was repeated over the three cropping seasons of 2003, 2004 and 2005. Irrespective of sites and intercrop design pea-barley intercropping improved the plant resource utilization (water, light, nutrients) to grain N yield with 25-30% using the Land Equivalent ratio. In terms of absolute quantities, sole cropped pea accumulated more N in the grains as compared to the additive design followed by the replacement design and then sole cropped barley. The post harvest soil mineral N content was unaffected by the preceding crops. Under the following winter wheat, the lowest mineral N content was generally found in early spring. Variation in soil mineral N content under the winter wheat between sites and seasons indicated a greater influence of regional climatic conditions and long-term cropping history than annual preceding crop and residue quality. Just as with the soil mineral N, the subsequent crop response to preceding crop was negligible. Soil N balances showed general negative values in the 2-year period, indicating depletion of N independent of preceding crop and cropping strategy. It is recommended to develop more rotational approaches to determine subsequent crop effects in organic cropping systems, since preceding crop effects, especially when including legumes, can occur over several years of cropping.

Heterotic and seed rate effects on nitrogen efficiencies in wheat

Four field experiments over 2 years investigated whether wheat hybrids had higher nitrogen-use efficiency (NUE) than their parents over a range of seed rates and different N regimes. There was little heterosis for total N in the above-ground biomass (NYt), but there was high-parent heterosis for grain N yields (NYg) in two of the hybrids, Hyno Esta and Hyno Rista, associated with greater nitrogen harvest index (NHI). Overall, the hybrids did not significantly increase the total dry matter produced per unit N in the above-ground crop (NUtE(t)), but did increase the grain dry matter per unit N in the above ground crop (NUtE(g)). The improvement in NUtE(g) was at the partial detriment of grain N concentration. Heterosis for grain NYg in Hyno Esta was lower at zero-N, suggesting that it did not achieve higher yields through more efficient capture or utilization of N. The greater NHI in Hyno Esta appeared to be facilitated by both greater N uptake, and remobilization of N from vegetative tissues, after anthesis. The response of N efficiency and uptake to seed rate was dependent on N supply and season. Where N fertilizer was applied, N uptake over time was slower at the lower seed rates, but where N was withheld N capture at the lowest seed rate soon approached the N capture of the higher seed rates. During grain filling, the rate of accumulation of N into the grain increased with seed rate and the duration of N accumulation decreased with seed rate. With N applied, N yields increased to all asymptote with seed rate, when N was withheld there was little response of N yields to seed rate. In 2002, N utilization efficiency (NUtE(t) and NUtE(g)) also increased asymptotically with seed rate, but in 2003 seed rate had little effect on N utilization efficiency. When nitrogen fertilizer had not been applied, NHI consistently decreased with increasing seed rate. The timing of N application made little difference to NUE, NY, or NUtE.

The effect of supplementing maize stover with cowpea (Vigna unguiculata) haulms on the intake and growth performance of Ethiopian sheep

This study compared the effect of supplementing maize stover (MS) with cowpea (Vigna unguiculata) haulms or commercial concentrate (CC) on feed intake, nutrient digestibility, live weight gain and carcass yield of male Ethiopian Highland sheep. Two cowpea genotypes, 12688 (forage) and IT96D-774 (dual-purpose), were used. A randomised block design was applied with groups of eight sheep, blocked by weight, allocated to one of six treatments; MS ad libitum either unsupplemented or supplemented daily with 150 or 300g dry matter (DM) of either cowpea or CC. MS contained more neutral detergent fibre (NDF), acid detergent fibre (ADF) and lignin than either cowpeas or CC Crude protein (CP) content of the forage-type cowpeas was higher than either dual-purpose or CC, while MS had the lowest CP content Relative to the negative control group, cowpea at either level significantly (P < 0.01) increased both MS intake and total NDF and lignin. Supplementation significantly (P < 0.01) increased nitrogen (N) intakes relative to the negative control, with N intake for CC and dual-purpose cowpea (high level) being similar to the intakes for cowpeas at 150g. N intake with the forage-type cowpea offered at higher levels was significantly (P < 0.01) greater than the other groups. No significant differences (P > 0.01) in MS intake were identified between cowpeas at either level or CC and, although intake level of CC increased, it did not differ significantly from the negative control group. Supplementation significantly (P < 0.01) improved average daily gain, with the negative control group losing weight over the experimental period, and increased final live weight, carcass cold weight and dressing percentage. Supplementation significantly improved the apparent digestibility of DM, organic matter and NDF, with no significant difference found between cowpeas at either level. N retention was negative for sheep offered only MS, but positive with all supplements, with cowpeas improving N retention to a greater extent than CC. Interestingly, N retention/N intake was higher with cowpeas offered at the lower level suggesting an improvement in utilisation efficiency. The results indicate that the supplementation of MS with cowpea enhanced ruminant production through improvements in digestibility and intake. Further, as production improvements associated with the two levels of supplementation did not differ significantly, it is suggested that where limited quantities of cowpea are available, it may be of greater nutritional benefit to offer smaller quantities over an increased number of animal days.

Splanchnic metabolism of nitrogenous compounds and urinary nitrogen excretion in steers fed alfalfa under conditions of increased absorption of ammonia and L-arginine supply across the portal-drained viscera

Effects of increased ammonia and/or arginine absorption across the portal-drained viscera (PDV) on net splanchnic (PDV and liver) metabolism of nitrogenous compounds and urinary N excretion were investigated in six cathetenzed Hereford x Angus steers (501 +/- 1 kg BW) fed a 75% alfalfa:25% (as-fed basis) corn-soybean meal diet (0.523 MJ of ME/[kg BW0.15.d]) every 2 h without (27.0 g of N/kg of dietary DM) and with 20 g of urea/kg of dietary DM (35.7 g of N/kg of dietary DM) in a split-plot design. Net splanchnic flux measurements were obtained immediately before beginning and ending a 72-h mesenteric vein infusion of L-arginine (15 mmol/h). For 3 d before and during arginine infusion, daily urine voided was measured and analyzed for N composition. Feeding urea increased PDV absorption (P < 0.01) and hepatic removal (P < 0.01) of ammonia N, accounting for 80% of increased hepatic urea N output (P < 0.01). Numerical increases in net hepatic removal of AA N could account for the remaining portion of increased hepatic urea N output. Arginine infusion increased hepatic arginine removal (P < 0.01) and hepatic urea N output (P < 0.03) and switched hepatic ornithine flux from net uptake to net output (P < 0.01), but numerical changes in net hepatic removal of ammonia and AA N could not account fully for the increase in hepatic urea N output. Increases in urine N excretion equaled quantities of N fed as urea or infused as arginine. Estimated salivary urea N excretion was not changed by either treatment. Urea cycle regulation occurs via a complex interaction of mechanisms and requires N sources other than ammonia, but the effect of increased ammonia absorption on hepatic catabolism of individual AA in the present study was not significant.

Nitrogen recycling through the gut and the nitrogen economy of ruminants: An asynchronous symbiosis

The extensive development of the ruminant forestomach sets apart their N economy from that of nonruminants in a number of respects. Extensive pregastric fermentation alters the profile of protein reaching the small intestine, largely through the transformation of nitrogenous compounds into microbial protein. This process is fueled primarily by carbohydrate fermentation and includes extensive recycling of N between the body and gut lumen pools. Nitrogen recycling occurs via blood and gut lumen exchanges of urea and NH3, as well as endogenous gut and secretory N entry into the gut lumen, and the subsequent digestion and absorption of microbial and endogenous protein. Factors controlling urea transfer to the gut from blood, including the contributions of urea transporters, remain equivocal. Ammonia produced by microbial degradation of urea and dietary and endogenous AA is utilized by microbial fermentation or absorbed and primarily converted to urea. Therefore, microbial growth and carbohydrate fermentation affect the extent of NH3 absorption and urea N recycling and excretion. The extensive recycling of N to the rumen represents an evolutionary advantage of the ruminant in terms of absorbable protein supply during periods of dietary protein deficiency, or asynchronous carbohydrate and protein supply, but incurs a cost of greater N intakes, especially in terms of excess N excretion. Efforts to improve the efficiency of N utilization in ruminants by synchronizing fermentable energy and N availability have generally met with limited success with regards to production responses. In contrast, imposing asynchrony through oscillating dietary protein concentration, or infrequent supplementation, surprisingly has not negatively affected production responses unless the frequency of supplementation is less than once every 3 d. In some cases, oscillation of dietary protein concentration has improved N retention compared with animals fed an equal amount of dietary protein on a daily basis. This may reflect benefits of Orn cycle adaptations and sustained recycling of urea to the gut. The microbial symbiosis of the ruminant is inherently adaptable to asynchronous N and energy supply. Recycling of urea to the gut buffers the effect of irregular dietary N supply such that intuitive benefits of rumen synchrony in terms of the efficiency of N utilization are typically not observed in practice.

Effects of triazole and strobilurin fungicide programmes, with and without late-season nitrogen fertiliser, on the baking quality of Malacca winter wheat

Field experiments were conducted over 3 years to study the effect of applying triazole and strobilurin fungicides on the bread-making quality of Malacca winter wheat. Averaged over all years the application of a fungicide programme increased yields, particularly when strobilurin fungicides were applied. Reductions in protein concentration, sulphur concentration, Hageberg failing number and loaf volumes also occurred as the amount of fungicide applied increased. However, there were no deleterious effects of fungicide application on sodium dodecyl sulphate (SDS) sedimentation volumes, N:S ratios or dough theology. Effects of fungicide application on bread-making quality were not product specific. Therefore, it appears that new mechanisms to explain strobilurin effects on bread-making quality do not need to be invoked. Where reductions in protein concentration did occur they could be compensated for by a late-season application of nitrogen either as granular ammonium nitrate at flag leaf emergence or foliar urea at anthesis. These applications, however, sometimes increased the N:S ratio of the extracted flour and failed to improve loaf volume. Multiple regression analysis revealed that main effects of year, flour protein concentration and N:S ratio could explain 93% of the variance in loaf volume caused by season, fungicide and nitrogen treatments. However, an equally good fit was achieved by just including sulphur concentration with year. (C) 2004 Elsevier Ltd. All rights reserved.

The effects of triazole and strobilurin fungicide programmes on nitrogen uptake, partitioning, remobilization and grain N accumulation in winter wheat cultivars

Field experiments were conducted over 3 years to assess the effect of a triazole fungicide programme, and additions of strobilurin fungicides to it, on nitrogen uptake, accumulation and partitioning in a range of winter wheat cultivars. Commensurate with delayed senescence, fungicide programmes, particularly when including strobilurins, improved grain yield through improvements in both crop biomass and harvest index, although the relationship with green area duration of the flag leaf (GFLAD) depended on year and in some cases, cultivar. In all years fungicide treatments significantly increased the amount of nitrogen in the above-ground biomass, the amount of nitrogen in the grain and the nitrogen harvest index. All these effects could be linearly related to the fungicide effect on GFLAD. These relationships occasionally interacted with cultivar but there was no evidence that fungicide mode of action affected the relationship between GFLAD and yield of nitrogen in the grain. Fungicide treatments significantly reduced the amount of soil mineral N at harvest and when severe disease had been controlled, the net remobilization of N from the vegetation to the grain after anthesis. Fungicide maintained the filling of grain with both dry matter and nitrogen. The proportionate accumulation of nitrogen in the grain was later than that of dry matter and this difference was greater when fungicide had been applied. Effects of fungicide on grain protein concentration and its relationship with GFLAD were inconsistent over year and cultivar. There were several instances where grain protein concentration was unaffected despite large (1(.)5 t/ha) increases in grain yield following fungicide use. Dilution of grain protein concentration following fungicide use, when it did occur, was small compared with what would be predicted by adoption of other yield increasing techniques such as the selection of high yielding cultivars (based on currently available cultivars) or by growing wheat in favourable climates.

The effects of adding picoxystrobin, azoxystrobin and nitrogen to a triazole programme on disease control, flag leaf senescence, yield and grain quality of winter wheat

The effect of adding strobilurins to a triazole (epoxiconazole) fungicide programme on the quality of a range of wheat cultivars was assessed in field experiments in three successive years. Strobilurin was applied at just flag leaf emergence (azoxystrobin) or at the start of stem extension (azoxystrobin or picoxystrobin) and again at flag leaf emergence or at flag leaf emergence and again at ear emergence (azoxystrobin). All strobilurin treatments reduced disease levels, delayed senescence of the flag leaf and consistently increased yields, thousand grain weight and specific weight. Reductions in Hagberg falling number were observed, even by fungicide applications at the start of stem extension, but effects were small compared to the variation among cultivars. Application of fungicide (triazole or strobilurin) before ear emergence increased the amount of blackpoint, but this was partly countered by applying azoxystrobin at ear emergence. The effect of fungicide on protein concentration differed over seasons and cultivar. Where they occurred. small reductions in protein concentration could be compensated for by extra application of nitrogen as foliar urea at anthesis. Foliar urea (40 kg N ha(-1)) applied at anthesis also improved Hagberg failing number and reduced blackpoint in one of the growing seasons. In one season, the effect of foliar urea at anthesis was compared with applications of granular fertiliser at flag leaf emergence. The granular treatment produced grain with more concentrated protein, while the later, foliar application produced higher specific weights. (C) 2003 Elsevier Science Ltd. All rights reserved.

Nitrogen budgets for three cropping systems fertilised with cattle manure

A field plot experiment was set up on a sandy loam soil of SW England in order to determine the efficiency of nitrogen use from different cattle manures. The manure treatments were low and high dry matter cattle slurries and one farmyard manure applied at a target rate of 200 kg total Nha(-1) year(-1), and an untreated control. There were three different cropping systems: ryegrass/clover mixture, maize/rye and maize/bare soil, which were evaluated during 1998/99 and 1999/00. Measurements were made of N losses, N uptake and herbage DM yields. Result showed that manure type had a significant effect on N utilisation only for maize. N balances were negative in maize (approximately -247 to -10 kg N) compared to grass (approximately 5-158 kg N). Agronomic management was more important than manure type in influencing N losses, where soil cultivation appeared to be a key factor when comparing maize and grass systems. (C) 2004 Elsevier Ltd. All rights reserved.

The influence of winter oilseed rape (Brassica napus ssp oleifera var. biennis) canopy size on grass weed growth and grass weed seed return

Four experiments conducted over three seasons (2002-05) at the Crops Research Unit, University of Reading, investigated effects of canopy management of autumn sown oilseed rape (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) on competition with grass weeds. Emphasis was placed on the effect of the crop on the weeds. Rape canopy size was manipulated using sowing date, seed rate and the application of autumn fertilizer. Lolium multiflorum Lam., L. x boucheanum Kunth and Alopecurus myosuroides Huds. were sown as indicative grass weeds. The effects of sowing date, seed rate and autumn nitrogen on crop competitive ability were correlated with rape biomass and fractional interception of photosynthetically active radiation (PAR) by the rape floral layer, to the extent that by spring there was good evidence of crop: weed replacement. An increase in seed rate up to the highest plant densities tested increased both rape biomass and competitiveness, e.g. in 2002/3, L. multiflorum head density was reduced from 539 to 245 heads/m(2) and spikelet density from 13 170 to 5960 spikelets/m(2) when rape plant density was increased from 16 to 81 plants/m(2). Spikelets/head of Lolium spp. was little affected by rape seed rate, but the length of heads of A. myosuroides was reduced by 9 % when plant density was increased from 29-51 plants/m(2). Autumn nitrogen increased rape biomass and reduced L. multiflorum head density (415 and 336 heads/m(2) without and with autumn nitrogen, respectively) and spikelet density (9990 and 8220 spikelets/m(2) without and with autumn nitrogen, respectively). The number of spikelets/head was not significantly affected by autumn nitrogen. Early sowing could increase biomass and competitiveness, but poor crop establishment sometimes overrode the effect. Where crop and weed establishment was similar for both sowing dates, a 2-week delay (i.e. early September to mid-September) increased L. multiflorum head density from 226 to 633 heads/m(2) and spikelet density from 5780 to 15 060 spikelets/m(2).

The effect of environment on endosperm cell-wall development in Triticum aestivum during grain filling: an infrared spectroscopic imaging study

One of the major factors contributing to the failure of new wheat varieties is seasonal variability in end-use quality. Consequently, it is important to produce varieties which are robust and stable over a range of environmental conditions. Recently developed sample preparation methods have allowed the application of FT-IR spectroscopic imaging methods to the analysis of wheat endosperm cell wall composition, allowing the spatial distribution of structural components to be determined without the limitations of conventional chemical analysis. The advantages of the methods, described in this paper, are that they determine the composition of endosperm cell walls in situ and with minimal modification during preparation. Two bread-making wheat cultivars, Spark and Rialto, were selected to determine the impact of environmental conditions on the cell-wall composition of the starchy endosperm of the developing and mature grain, focusing on the period of grain filling (starting at about 14 days after anthesis). Studies carried out over two successive seasons show that the structure of the arabinoxylans in the endosperm cell walls changes from a highly branched form to a less branched form. Furthermore, during development the rate of restructuring was faster when the plants were grown at higher temperature with restricted water availability from 14 days after anthesis with differences in the rate of restructuring occurring between the two cultivars.

The effect of acetylsalicylic acid and oxalic acid on Myzus persicae and Aphidius colemani

Plants can respond to damage by pests with both induced direct defences and indirect defences by the attraction of their natural enemies. Foliar application of several plant-derived chemicals, such as salicylic acid and oxalic acid, can induce these defence mechanisms. The effect of acetylsalicylic acid and oxalic acid on the aphid Myzus persicae Sulzer (Homoptera: Aphididae) and its parasitoid Aphidius colemani Viereck (Hymenoptera: Aphidiidae) was investigated. Experiments were carried out with direct application of acetylsalicylic and oxalic acids on these insects, as well as choice and no-choice tests using foliar application of both chemicals on Brussels sprouts plants, Brassica oleracea var. gemmifera L. (Brassicaceae). Parasitoids were given a choice between treated and untreated plants for oviposition, and the effects of the chemicals on aphid and parasitoid development were determined. Although direct application of both chemicals increased aphid mortality, their foliar application did not induce resistance against aphids. The foliar application of such compounds, even in low concentration as shown in the choice tests, has the potential to induce indirect plant defences against aphids by encouraging aphid parasitisation. Although the direct application of both chemicals reduced parasitoid emergence from their hosts, the foliar application of acetylsalicylic acid and low concentrations of oxalic acid did not have a negative effect on parasitoid emergence ability. However, 10 mm oxalic acid reduced the number of emerged parasitoids in no-choice experiments. This study shows that foliar application of acetylsalicylic and oxalic acids has the potential to encourage aphid parasitisation, but care is needed as high concentrations of oxalic acid can have a negative effect on these beneficial organisms.

Transcriptomic analysis of rhizobium leguminosarum biovar viciae in symbiosis with host plants pisum sativum and Vicia cracca

Rhizobium leguminosarum bv. viciae forms nitrogen-fixing nodules on several legumes, including pea (Pisum sativum) and vetch (Vicia cracca), and has been widely used as a model to study nodule biochemistry. To understand the complex biochemical and developmental changes undergone by R. leguminosarum bv. viciae during bacteroid development, microarray experiments were first performed with cultured bacteria grown on a variety of carbon substrates (glucose, pyruvate, succinate, inositol, acetate, and acetoacetate) and then compared to bacteroids. Bacteroid metabolism is essentially that of dicarboxylate-grown cells (i.e., induction of dicarboxylate transport, gluconeogenesis and alanine synthesis, and repression of sugar utilization). The decarboxylating arm of the tricarboxylic acid cycle is highly induced, as is gamma-aminobutyrate metabolism, particularly in bacteroids from early (7-day) nodules. To investigate bacteroid development, gene expression in bacteroids was analyzed at 7, 15, and 21 days postinoculation of peas. This revealed that bacterial rRNA isolated from pea, but not vetch, is extensively processed in mature bacteroids. In early development (7 days), there were large changes in the expression of regulators, exported and cell surface molecules, multidrug exporters, and heat and cold shock proteins. fix genes were induced early but continued to increase in mature bacteroids, while nif genes were induced strongly in older bacteroids. Mutation of 37 genes that were strongly upregulated in mature bacteroids revealed that none were essential for nitrogen fixation. However, screening of 3,072 mini-Tn5 mutants on peas revealed previously uncharacterized genes essential for nitrogen fixation. These encoded a potential magnesium transporter, an AAA domain protein, and proteins involved in cytochrome synthesis.

Legumes regulate Rhizobium bacteroid development and persistence by the supply of branched-chain amino acids

One of the largest contributions to biologically available nitrogen comes from the reduction of N-2 to ammonia by rhizobia in symbiosis with legumes. Plants supply dicarboxylic acids as a carbon source to bacteroids, and in return they receive ammonia. However, metabolic exchange must be more complex, because effective N-2 fixation by Rhizobium leguminosarum bv viciae bacteroids requires either one of two broad-specificity amino acid ABC transporters (Aap and Bra). It was proposed that amino acids cycle between plant and bacteroids, but the model was unconstrained because of the broad solute specificity of Aap and Bra. Here, we constrain the specificity of Bra and ectopically express heterologous transporters to demonstrate that branched-chain amino acid (LIV) transport is essential for effective N-2 fixation. This dependence of bacteroids on the plant for LIV is not due to their known down-regulation of glutamate synthesis, because ectopic expression of glutamate dehydrogenase did not rescue effective N-2 fixation. Instead, the effect is specific to LIV and is accompanied by a major reduction in transcription and activity of LIV biosynthetic enzymes. Bacteroids become symbiotic auxotrophs for LIV and depend on the plant for their supply. Bacteroids with aap bra null mutations are reduced in number, smaller, and have a lower DNA content than wild type. Plants control LIV supply to bacteroids, regulating their development and persistence. This makes it a critical control point for regulation of symbiosis. MICROBIOLOGY