A temporal limit to the association between flag leaf life extension by fungicides and wheat yields

Three field experiments, each repeated over two or three seasons, on winter wheat investigated a possible limit to the association between grain yield and flag leaf life, as extended by fungicide application. The experiments involved up to six cultivars and different application rates, timings and frequencies of the strobilurin azoxystrobin and the triazole epoxiconazole. In the 2000/01 and 2001/02 seasons, the relationships between the thermal time to 37 % green flag leaf area (m) and yield deviated from linearity. 'Broken stick' models were fitted to cultivar x experiment combinations within each season and the limit to the benefit to yield associated with extending flag leaf life was 700 degrees C days (S.E. = 20.7) and 725 degrees C days (S.E. = 9.33) after anthesis in 2000/01 and 2001/02, respectively. In 2002/03, the relationship between yield and in did not deviate significantly (P > 0.05) from linearity, but in this latter year the fungicide application failed to increase In past 700 degrees C days. (c) 2004 Elsevier B.V. All rights reserved.

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.

Mapping quantitative trait loci for flag leave senescence as a yield determinant in winter wheat under optimal and drought stressed environments

The timing of flag leaf senescence (FLS) is an important determinant of yield under stress and optimal environments. A doubled haploid population derived from crossing the photo period-sensitive variety Beaver,with the photo period-insensitive variety Soissons, varied significantly for this trait, measured as the percent green flag leaf area remaining at 14 days and 35 days after anthesis. This trait also showed a significantly positive correlation with yield under variable environmental regimes. QTL analysis based on a genetic map derived from 48 doubled haploid lines using amplified fragment length polymorphism (AFLP) and simple sequence repeat (SSR) markers, revealed the genetic control of this trait. The coincidence of QTL for senescence on chromosomes 2B and 2D under drought-stressed and optimal environments, respectively, indicate a complex genetic mechanism of this trait involving the re-mobilisation of resources from the source to the sink during senescence.

Genotype and fungicide effects on late-season root growth of winter wheat

The aim of this work was to investigate differences among genotypes in post-anthesis root growth and distribution of modern UK winter wheat cultivars, and the effects of fungicide applications. Post-anthesis root growth of up to six cultivars of winter wheat (Triticum aestivum L.), given either one or three applications of fungicide, was studied in field experiments during two seasons. Total root mass remained unchanged between GS63 (anthesis) and GS85, but root length increased significantly from 14.7 to 31.4 km m(2) in one season. Overall, there was no evidence for a decline in either root mass or length during grain filling. Root mass as a proportion of total plant mass was about 0.05 at GS85. There were significant differences among cultivars in root length and mass especially below 30 cm. Malacca had the smallest root length and Savannah the largest, and Shamrock had a significantly larger root system below 40 cm in both seasons. Fungicide applied at ear emergence had no significant effect on root mass in either season but increased root length (P < 0.01) in the more disease-prone season. By maintaining a green canopy for longer, fungicide applied at flag leaf emergence may have resulted in delayed senescence of the root system and contributed to the post-anthesis maintenance of root mass and length.

Influence of foliar diseases and their control by fungicides on grain yield and quality in wheat

Twenty-eight field experiments on sandy-loam soils in the UK (1982-2003) are reviewed by relating the extension of the green area duration of the flag leaf (GLADF) by fungicides to effects on yield and quality of winter wheat. Over all experiments mean grain yield = 8.85t ha(-1) at 85% DM. With regards quality, mean values were: thousand grain weight (TGW) = 44.5 g; specific weight (SWT) = 76.9 kg hl(-1); crude protein concentration (CP (N x 5.7)) = 12.5 % DM; Hagberg falling number (HFN) = 285 s; and sodium dodecyl sulphate (SDS)-sedimentation volume = 69ml. For each day (d) that fungicides increased GLADF there were associated average increases in yield (0.144 1 ha(-1) d(-1), se 0.0049, df = 333), TGW (0.56 gd(-1), se = 0.017) and SWT (0.22 kg hl(-1) d(-1), se 0.011). Some curvature was evident in all these relationships. When GLADF was delayed beyond 700 degrees Cd after anthesis, as was possible in cool wet seasons, responses were curtailed, or less reliable. Despite this apparent terminal sink limitation, fungicide effects on sink size, eg endosperm cell numbers or maximum water mass per grain, were not prerequisites for large effects on grain yield, TGW or SWT. Fungicide effects on CP were variable. Although the average response of CP was negative (-0.029%DM/d; se = 0.00338), this depended on cultivar and disease controlled. Controlling biotrophs such as rusts, (Puccinia spp.) tended to increase CP, whereas controlling a more necrotrophic pathogen (Septoria tritici) usually reducedCP. Irrespective of pathogen controlled, delaying senescence of the flag leaf was associated with increased nitrogen yields in the grain (averaging 2.24 kg N ha-1 d(-1), se = 0.0848) due to both increased N uptake into the above ground crop, and also more efficient remobilisation of N from leaf laminas. When sulphur availability appeared to be adequate, fungicide x cultivar interactions were similar on S as for CP, although N:S ratios tended to decline (i.e. improve for bread making) when S. tritici was controlled. On average, SDS-sedimentation volume declined (-0. 18 ml/d, se = 0.027) with increased GLADF, broadly commensurate with the average effect on CP. Hagberg falling number decreased as fungicide increased GLADF (-2.73 s/d, se = 0.178), indicating an increase in alpha-amylase activity.

Fungicide and cultivar affect post-anthesis patterns of nitrogen uptake, remobilization and utilization efficiency in wheat

Three successive field experiments (2000/01-2002/03) assessed the effect of wheat cultivar (Consort.. Hereward and Shamrock) and fungicide (epoxiconazole and azoxystrobin) applied at and after flag leaf emergence on the nitrogen in the above-ground crop (Total N) and grain (Grain N), net nitrogen remobilization from non-grain tissues (Remobilized N). grain dry matter (Grain Dill), and nitrogen utilization efficiency (NUtE(g) = Grain DM/Total N). Ordinary logistic curves were fitted to the accumulation of Grain N, Grain DM and Remobilized N against thermal time after anthesis and used to simultaneously derive fits for Total N and NUtE(g). When disease was controlled, Consort achieved the greatest Grain DM, Total N, Grain N and NUtEg; in each case due mostly to longer durations, rather than quicker rates, of accumulation. Fungicide application increased final Grain Dill.. Grant N, Total N and Remobilized N, also mostly through effects on duration rather than rate of accumulation. Completely senesced leaf laminas retained less nitrogen when fungicide had been applied compared with leaf laminas previously infected severely with brown rust (Puccinia recondita) and Septoria tritici, or with just S. tritici. Late movement of nitrogen out of fungicide-treated laminas contributed to extended duration of both nitrogen remobilization and grain N filling, and meant that increases in NUtE(g) could occur without simultaneous reductions in grain N concentration.

Modelling simultaneously water content and dry matter dynamics of wheat grains

A model was devised to describe simultaneously the grain masses of water and dry matter against thermal time during grain filling and maturation of winter wheat. The model accounted for a linear increase in water mass of duration anthesis-m(1) (end of rapid water assimilation phase) and rate a, followed by a more stable water mass until in,, after which water mass declined rapidly at rate e. Grain dry matter was described as a linear increase of rate bgf until a maximum size (maxgf) was attained at m(2).The model was fitted to plot data from weekly samples of grains taken from replicated field experiments investigating effects of grain position (apical or medial), fungicide (five contrasting treatments), sowing date (early or late), cultivar (Malacca or Shamrock) and season (2001/2002 and 2002/2003) on grain filling. The model accounted for between 83 and 99% of the variation ( 2) when fitted to data from individual plots, and between 97 and 99% when fitted to treatment means. Endosperm cell number of grains from early-sown plots in the first season were also counted. Differences in maxgf between grain positions and also between cultivars were mostly the result of effects on bgf and were empirically associated with water mass at nil. Fungicide application controlled S. tritici and powdery mildew infection, delayed flag leaf senescence, increased water mass at m(1) (wm(1)), and also increased m(2), bgf and maxgf. Fungicide effects on water mass were detected before fungicide effects on dry matter, but comparison of the effects of individual fungicide treatments showed no evidence that effects on wm(1), nor on endosperm cell numbers at about m(1), were required for fungicide effects on maxgf, (c) 2005 Elsevier B.V. All rights reserved.

Delaying senescence of wheat with fungicides has interacting effects with cultivar on grain sulphur concentration but not with sulphur yield or nitrogen: sulphur ratios

Winter wheat was grown in three field experiments, each repeated over two or three seasons, to investigate effects of extending flag leaf life by fungicide application on the concentration, kg ha(-1) and mg grain(-1) of nitrogen (N) and sulphur (S) as well as N:S ratio and sodium dodecyl sulphate (SDS) sedimentation volume. The experiments involved up to six cultivars and different application rates, timings and frequencies of azoxystrobin and epoxiconazole. For every day the duration to 37 % green flag leaf area (m) was extended, N yield was increased by 2.58 kg ha(-1), N per grain by 0.00957 mg, S yield by 0.186 kg ha(-1) and S per grain by 0.000718 mg. The N:S ratio decreased by 0.0135 per day. There was no evidence that these responses varied with cultivar. In contrast, the relationship between flag leaf life and N or S concentration interacted with cultivar. The N and S concentrations of Shamrock, the cultivar that suffered most from brown rust (Puccinia rccondita), increased with the extension of flag leaf life whereas the concentrations of N and S in Malacca, a cultivar more susceptible to Septoria tritici, decreased as flag leaf senescence was delayed. This was because the relationships between m and N and S yields were much better conserved over cultivars than those between m and thousand grain weight (TGW) and grain yield ha(-1). (c) 2004 Elsevier B.V. All rights reserved.

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.

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.

Apical leaf necrosis and leaf nitrogen dynamics in diseased leaves: a model study

Apical leaf necrosis is a physiological process related to nitrogen (N) dynamics in the leaf. Pathogens use leaf nutrients and can thus accelerate this physiological apical necrosis. This process differs from necrosis occurring around pathogen lesions (lesion-induced necrosis), which is a direct result of the interaction between pathogen hyphae and leaf cells. This paper primarily concentrates on apical necrosis, only incorporating lesion-induced necrosis by necessity. The relationship between pathogen dynamics and physiological apical leaf necrosis is modelled through leaf nitrogen dynamics. The specific case of Puccinia triticina infections on Triticum aestivum flag leaves is studied. In the model, conversion of indirectly available N in the form of, for example, leaf cell proteins (N-2(t)) into directly available N (N-1(t), i.e. the form of N that can directly be used by either pathogen or plant sinks) results in apical necrosis. The model reproduces observed trends of disease severity, apical necrosis and green leaf area (GLA) and leaf N dynamics of uninfected and infected leaves. Decreasing the initial amount of directly available N results in earlier necrosis onset and longer necrosis duration. Decreasing the initial amount of indirectly available N, has no effect on necrosis onset and shortens necrosis duration. The model could be used to develop hypotheses on how the disease-GLA relation affects yield loss, which can be tested experimentally. Upon incorporation into crop simulation models, the model might provide a tool to more accurately estimate crop yield and effects of disease management strategies in crops sensitive to fungal pathogens.

Temporal patterns of nutrient availability around nests of leaf-cutting ants (Atta colombica) in secondary moist tropical forest

Leaf-cutting ants consume up to 10% of canopy leaves in the foraging area of their colony and therefore represent a key perturbation in the nutrient cycle of tropical forests. We used a chronosequence of nest sites on Barro, Colorado Island, Panama, to assess the influence of leaf-cutting ants (Atta colombica) on nutrient availability in a neotropical rainforest. Twelve nest sites were sampled, including active nests, recently abandoned nests (<1 year) and long-abandoned nests (>1 year). Waste material discarded by the ants down-slope from the nests contained large concentrations of nitrogen and phosphorus in both total and soluble forms, but decomposed within one year after the nests were abandoned. Despite this, soil under the waste material contained high concentrations of nitrate and ammonium that persisted after the disappearance of the waste, although soluble phosphate returned to background concentrations within one year of nest abandonment. Fine roots were more abundant in soil under waste than control soils up to one year after nest abandonment, but were not significantly different for older sites. In contrast to the waste dumps, soil above the underground nest chambers consistently contained lower nutrient concentrations than control soils, although this was not statistically significant. We conclude that the 'islands of fertility' created by leaf-cutting ants provide a nutritional benefit to nearby plants for less than one year after nest abandonment in the moist tropical environment of Barro Colorado Island. Published by Elsevier Ltd.

Global transcript analysis of rice leaf and seed using SAGE technology.

We have compiled two comprehensive gene expression profiles from mature leaf and immature seed tissue of rice (Oryza sativa ssp. japonica cultivar Nipponbare) using Serial Analysis of Gene Expression (SAGE) technology. Analysis revealed a total of 50 519 SAGE tags, corresponding to 15 131 unique transcripts. Of these, the large majority (approximately 70%) occur only once in both libraries. Unexpectedly, the most abundant transcript (approximately 3% of the total) in the leaf library was derived from a type 3 metallothionein gene. The overall frequency profiles of the abundant tag species from both tissues differ greatly and reveal seed tissue as exhibiting a non-typical pattern of gene expression characterized by an over abundance of a small number of transcripts coding for storage proteins. A high proportion ( approximately 80%) of the abundant tags (> or = 9) matched entries in our reference rice EST database, with many fewer matches for low abundant tags. Singleton transcripts that are common to both tissues were collated to generate a summary of low abundant transcripts that are expressed constitutively in rice tissues. Finally and most surprisingly, a significant number of tags were found to code for antisense transcripts, a finding that suggests a novel mechanism of gene regulation, and may have implications for the use of antisense constructs in transgenic technology.