Influence of direct-drilling of winter wheat on incidence of take-all and eyespot

Winter wheat drilled directly into stubble or pasture treated with paraquat to kill the vegetation has been found to be less severely attacked by take-all (Ophiobolus graminis (Sacc.) Sacc.) and eyespot (Cercosporella herpotrichoides Fron) than wheat drilled after cultivation. The reduction of take-all is associated, not with a direct effect of the chemical, but with factors, resulting from the technique, which limit the rate of spread of the fungus in the undisturbed soil.     

Experiments on soil drenching with fungicides against take-all in wheat

In short term pot experiments benomyl, iprodione and KWG 0599 applied as soil drenches in several types of soil significantly suppressed take-all symptoms from inoculum placed just below wheat seeds planted 1×5 cm deep, and in sand but not other soils when seeds were 5 cm deep. Benomyl was, however, effective against inoculum below seed planted 5 cm deep in a loam-sand mixture when the drench contained an alcohol ethoxylate surfactant. Computer simulations of fungicide distributions in the soils correlated well with disease control observations. In long term outdoor pot experiments two drenches with benomyl (without surfactant) controlled disease significantly for at least 3 months against inoculum placed 15 cm deep. The significance of these results for the practical control of take-all by fungicides is discussed.     

Effects of eyespot on the yield of winter wheat

In two experiments, on Julia and Zephyr spring barley, evidence was obtained that the development of powdery mildew in sprayed plots depended very much on their proximity to unsprayed crop. The effective period of control achieved with one spray was greater where plots were well separated from unsprayed crop than where they were surrounded by or very close to unsprayed crop. There is probably a risk that small plot experiments may not only underestimate the potential effectiveness of fungicides in agricultural practice, but also give misleading information on optimal spray timing.     

Prospects for fungicidal control of take-all of wheat

A range of fungicides and herbicides was tested against Gaeumannomyces graminis (causal agent of take-all) on agar plates, and on wheat seedlings in pots and in liquid culture. Benomyl, the standard in all tests, was consistently most effective: like iprodione, nuarimol and KWG 0599 , it diminished infection from inoculum placed just below the seeds more effectively when applied as a drench than as a soil-mix. Benomyl as a soil-mix was most effective in soils with least organic matter. Some compounds toxic to the pathogen on agar plates and in plants grown in liquid culture were ineffective as soil treatments. The practical limitations of soil treatment with conventional fungicides and application methods are discussed.     

Role of antibiotics and siderophores in biocontrol of take-all disease of wheat

Both antibiotics and siderophores have been implicated in the control of soilborne plant pathogens by fluorescent pseudomonads. In Pseudomonas fluorescens 2–79, which suppresses take-all of wheat, the importance of the antibiotic phenazine-1-carboxylic acid was established with mutants deficient or complemented for antiobiotic production and by isolation of the antibiotic from the roots of wheat colonized by the bacteria. Genetic and biochemical studies of phenazine synthesis have focused on two loci; the first is involved in production of both anthranilic acid and phenazine-1-carboxylic acid, and the second encodes genes involved directly in phenazine synthesis. Because the antibiotic does not account fully for the suppressiveness of strain 2-79, additional mutants were analyzed to evaluate the role of the fluorescent siderophore and of an antifungal factor (Aff, identified as anthranilic acid) that accumulates when iron is limiting. Whereas strains producing only the siderophore conferred little protection against take-all, Aff⁺ strains were suppressive, but much less so than phenazine-producing strains. Iron-regulated nonsiderophore antibiotics may be produced by fluorescent pseudomonads more frequently than previously recognized, and could be partly responsible for beneficial effects that were attributed in the past to fluorescent siderophores.     

Effects of water potential and temperature on the development of eyespot lesions in wheat

Growth of Pseudocercosporella herpotrichoides on potato dextrose agar at water potentials from -0.5 to -6.9 MPa was optimal at 20°C. At 12 and 20°C, six isolates of P. herpotrichoides grew more rapidly at -0.5 to -2 MPa than at -6.9 MPa. Wheat plants inoculated with P. herpotrichoides and grown in columns of soil at either 15 or 20°C developed more severe eyespot lesions under a heavy watering regime than under medium or light watering regimes. P. herpotrichoides penetrated leaf sheaths of inoculated plants grown in compost more rapidly at 10°C night/15°C day temperatures than at 5/10°C; death of leaf sheaths was also more rapid and consequently there were fewer living infected leaf sheaths at 10/15°C than at 5/10°C. Irrigating for 5 wk before harvest increased the severity of eyespot lesions in a 1983 wheat crop.     

Effects of ammonium and nitrate nutrition or take-all disease of wheat in pots

Plant debris, naturaiiy infested with the take-all fungus (Ophiobolus graminis), was washed from soil and added to a leached sandy loam, deficient in nitrate nitrogen (NO₃-N) and magnesium. Nutrient solutions containing potassium and phosphorus, with and without magnesium, were added to the amended soil unsupplemented, or with either NO₃-N, ammonium nitrogen (NH₄+-N), or both. Nitrification of NH₄+-N was inhibited by 2–chloro-6–(trichloromethy1)-pyridine (N-Serve). After 38 days at 19°C, fewer plants had take-all with N (75 or 100 mg/kg soil) than without and root systems were most discoloured and had most diseased axes when nutrients were not added. Plants given NH₄+-N developed less take-all when magnesium was present. A comparison of forms of N in the presence of added magnesium showed that take-all was least with a mixture of both forms of N, intermediate with NO₃-N alone and worst with NH₄+-N alone. The most extensive lesions on individual root axes occurred on plants given NH₄+-N. It is suggested that take-all will be least when the amounts and ratio of NH₄+-N and NO₃-N are optimum for the growth of the host.     

Nitrate and ammonium in the rhizosphere of wheat crops and concurrent observations of take-all

Summary In 1973 the concentration of nitrate nitrogen in the rhizosphere of winter wheat (NO₃-N_r, extracted withN-K₂SO₄ from soil from within 5 mm of the roots) growing in slightly acid, flintly loam decreased progressively from January until early April, when it became less than that of ammonium nitrogen in the rhizosphere (NH₄-N_r, extracted withN-K₂SO₄). An April application of ammonium nitrate (as Nitro-chalk) augmented NO₃-N_rmore than NH₄-N_r, but because NH₄-N_rwas depleted less quickly it exceeded NO₃-N_rfrom early June onwards. From January to April pH_rremained between 6.9–7.2, but then decreased progressively to a minimum of 6.3 in June. The differences in soil pH and forms of N amongst plots that had grown three, six or fourteen consecutive cereal crops susceptible to take-all were small; a first wheat crop differed only in having less extreme changes in NO₃-N_rduring April-June.First winter wheat crops were grown on a similar soil in 1974–5, either with farmyard manure applied in autumn (FYM plot, bulk soil pH 7.4) or Nitro-chalk applied in April (NPK plot, pH_b 6.7). Soil nitrate decreased considerably in May and to a lesser extent in June and was always at a lower concentration in the rhizosphere. NH₄-N_rwas more than NO₃-N_rin water extracts after May in the FYM plot and after June in the NPK plot, but NH₄-N_bnever exceeded NO₃-N_b.There was, therefore, a tendency for more of the dwindling mineral N to exist as NH₄-N close to roots in spring and summer unless temporarily disrupted by spring fertilizer. Take-all developed against this background was mostly slight and never severe. In 1973 it was least in the first wheat, and most in the third, but the differences were small and disease increased steadily in all plots after mid-February. In 1975 it developed quickest and most in the FYM plot. re]19760928     

Soil-applied fungicides for controlling take-all in field experiments with winter wheat

Soil treatment fungicides were tested against take-all (Gaeumannomyces graminis var. tritici) in three field experiments with winter wheat. Fungicides were applied as drenches either before sowing in autumn, and incorporated by rotary harrowing, or to the crop in spring. The most effective treatments were autumn applied benomyl (20 kg/ha) and nuarimol (0·55-4·4 kg/ha). However, the highest nuarimol concentration depressed yield. Benomyl sometimes induced a resurgence of take-all in the second wheat crop after treatment. Nuarimol had no adverse effects in subsequent crops, and neither fungicide hindered the onset of take-all decline in a third crop after treatment. The possible value of soil treatment in future control strategies is discussed.     

Mapping QTL for resistance to eyespot of wheat in Aegilops longissima

Eyespot is an economically important disease of wheat caused by the soilborne fungi Oculimacula yallundae and O. acuformis. These pathogens infect and colonize the stem base, which results in lodging of diseased plants and reduced grain yield. Disease resistant cultivars are the most desirable control method, but resistance genes are limited in the wheat gene pool. Some accessions of the wheat wild relative Aegilops longissima are resistant to eyespot, but nothing is known about the genetic control of resistance. A recombinant inbred line population was developed from the cross PI 542196 (R) × PI 330486 (S) to map the resistance genes and better understand resistance in Ae. longissima. A genetic linkage map of the S(l) genome was constructed with 169 wheat microsatellite markers covering 1261.3 cM in 7 groups. F(5) lines (189) were tested for reaction to O. yallundae and four QTL were detected in chromosomes 1S(l), 3S(l), 5S(l), and 7S(l). These QTL explained 44 % of the total phenotypic variation in reaction to eyespot based on GUS scores and 63 % for visual disease ratings. These results demonstrate that genetic control of O. yallundae resistance in Ae. longissima is polygenic. This is the first report of multiple QTL conferring resistance to eyespot in Ae. longissima. Markers cfd6, wmc597, wmc415, and cfd2 are tightly linked to Q.Pch.wsu-1S ( l ), Q.Pch.wsu-3S ( l ), Q.Pch.wsu-5S ( l ), and Q.Pch.wsu-7S ( l ), respectively. These markers may be useful in marker-assisted selection for transferring resistance genes to wheat to increase the effectiveness of resistance and broaden the genetic diversity of eyespot resistance.     

Effects of wheat volunteers and blackgrass in set-aside following a winter wheat crop on soil infectivity and soil conduciveness to take-all

Two experiments were carried out in France in which disease indices were used to evaluate the effects of wheat volunteers and blackgrass (Alopecurus myosuroides) on soil infectivity and soil conduciveness to take-all caused by Gaeumannomyces graminis var. tritici. Soil infectivity was evaluated by measuring the disease index on susceptible wheat plants grown on soil samples collected from the field. Soil conduciveness to the disease was obtained by measuring disease indices on plants grown on soil samples to which different amounts of take-all fungus inoculum were added. One experiment (Expt. 1) was carried out using soils from farmers' fields (two fields in 1994 and two in 1995); soil infectivity and soil conduciveness were evaluated for three experimental situations: bare soil, soil with wheat volunteers and soil with blackgrass plants. In 1994 the soil infectivity was zero in bare soil, high with the wheat cover, and intermediate with the blackgrass cover. In 1995 the soil infectivity was uniformly low for all three conditions. Soils bearing wheat were less conducive than bare soil, soils bearing blackgrass and bare soils were similarly conducive. A second experiment (Expt. 2) carried out in 1995 compared the soil infectivity and soil conduciveness to take-all of soils planted with wheat or blackgrass in set-aside land after periods of wheat monoculture of 0–6 yr. The soil infectivity was low for all treatments. The soil was more conducive after blackgrass than after wheat. In both cases, the soil conduciveness was less when the monoculture had continued for more than 4 yr. The decline was less after blackgrass than after wheat. Thus, whenever set-aside is set up during the increase phase of the disease in fields with cereal successions, abundant wheat volunteers might hinder the expected positive effect of a break in cereal successions on take-all development. The presence of blackgrass in a set-aside field, with significant soil infectivity and high soil conduciveness, might increase the risks of take-all development in a wheat crop following set-aside.     

Changes in populations of rhizosphere bacteria associated with take-all disease of wheat

Take-all, caused by Gaeumannomyces graminis var. tritici, is one of the most important fungal diseases of wheat worldwide. Knowing that microbe-based suppression of the disease occurs in monoculture wheat fields following severe outbreaks of take-all, we analyzed the changes in rhizosphere bacterial communities following infection by the take-all pathogen. Several bacterial populations were more abundant on diseased plants than on healthy plants, as indicated by higher counts on a Pseudomonas-selective medium and a higher fluorescence signal in terminal restriction fragment length polymorphism analyses of amplified 16S ribosomal DNA (rDNA). Amplified rDNA restriction analysis (ARDRA) of the most abundant cultured populations showed a shift in dominance from Pseudomonas to Chryseobacterium species in the rhizosphere of diseased plants. Fluorescence-tagged ARDRA of uncultured rhizosphere washes revealed an increase in ribotypes corresponding to several bacterial genera, including those subsequently identified by partial 16S sequencing as belonging to species of alpha-, beta-, and gamma-proteobacteria, sphingobacteria, and flavobacteria. The functional significance of some of these populations was investigated in vitro. Of those isolated, only a small subset of the most abundant Pseudomonas spp. and a phlD(+) Pseudomonas sp. showed any significant ability to inhibit G. graminis var. tritici directly. When cultured strains were mixed with the inhibitory phlD(+) Pseudomonas strain, the Chryseobacterium isolates showed the least capacity to inhibit this antagonist of the pathogen, indicating that increases in Chryseobacterium populations may facilitate the suppression of take-all by 2,4-diacetylphloroglucinol-producing phlD(+) pseudomonads.     

A role for managanese in the resistance of wheat plants to take-all

Summary The hypothesis that wheat plants deficient in managenese are predisposed to infection byGaeumannomyces graminis is outlined, and a test of the hypothesis in a soil system is reported. The results supported the hypothesis: wheat plants growing in managanese-deficient soil, although not showing foliar symptoms, were markedly more susceptible to infection; plant analysis confirmed the nutrient status of the plants. A review of the literature on take-all in wheat coupled with the results of our experiments suggests a reinterpretation of the etiology of this disease, since those edaphic factors which promote infection by this organism are those which also render managese unavailable to the host. Managenese nutrition is proposed as a common factor in many of the environmental conditions which influence the host-pathogen balance.     

Field tests of bacteria and soil-applied fungicides as control agents for take-all in winter wheat

Putative biological and chemical treatments for controlling take-all were used in each of three consecutive years at two locations where winter wheat crops were grown in naturally-infested fields. The chemical treatments more often decreased take-all than the biological treatments, but no treatment consistently and significantly decreased take-all, nor did any cause a significant increase in yield. An isolate of Bacillus cereus var. mycoides and one of B. pumilis, applied as soil drenches in autumn or spring, or in the seed furrows, were usually ineffective. Of the few significant effects on disease, half were associated with increases and half with decreases, and most occurred in April and did not persist to late June. Two strains of Pseudomonas pluorescens applied to the seed were ineffective. The fungicide benomyl, applied as a drench in autumn and spring at 20 kg/ha was ineffective, while nuarimol, applied as a drench in autumn at 2 kg/ha was sometimes effective. Nuarimol incorporated into the seed bed at 2 kg/ha was the most effective treatment. In analyses using a functional relationship model for data from treated and untreated plots 12% of 176 data sets for biological treatments, 38% of 96 data sets for chemical treatments and 81% of 16 data sets for combined treatments showed increasing efficiency of the treatment with increasing disease intensity. These findings also demonstrate an additional advantage of the experimental design, namely that treatments are tested at different disease intensity levels within fields.     

Plant genotype, micronutrient fertilization and take-all infection influence bacterial populations in the rhizosphere of wheat

The relationship between micronutrient efficiency of four wheat (Triticum aestivum L.) genotypes, tolerance to take-all disease (caused by Gaeumannomyces graminis (Sacc.) Arx and Olivier var. tritici Walker), and bacterial populations in the rhizosphere was tested in soil fertilized differentially with Zn and Mn. Plant growth was reduced by Mn or Zn deficiency and also by take-all. There was an inverse relationship between micronutrient efficiency of wheat genotypes when grown in deficient soils and the length of take-all lesions on roots (efficient genotypes had shorter lesions than inefficient ones). In comparison to the rhizosphere of control plants of genotypes Aroona and C8MM receiving sufficient Mn and Zn, the total numbers of bacterial cfu (colony forming units) were greater in the rhizosphere of Zn-efficient genotype Aroona under Zn deficiency and in Mn-efficient genotype C8MM under Mn deficiency. These effects were not observed in other genotypes. Take-all decreased the number of bacterial cfu in the rhizosphere of fully-fertilized plants but not of those subjected to either Mn or Zn deficiency. In contrast, the Zn deficiency treatment acted synergistically with take-all to increase the number of fluorescent pseudomonads in the rhizosphere. Although numbers of Mn-oxidising and Mn-reducing bacteria were generally low, take-all disease increased the number of Mn reducers in the rhizosphere of Mn-efficient genotypes Aroona and C8MM. Under Mn-deficiency conditions, the number of Mn reducers in the rhizosphere increased in Aroona but not in C8MM wheat. The results suggest that bacterial microflora may play a role in the expression of Mn and Zn efficiency and tolerance to take-all in some wheat genotypes.     

Soil conditions and the take-all disease of wheat: VIII. Further experiments on the survival of Ophiobolus graminis in infected wheat stubble

Assimilable nitrogen in various forms prolonged the life of Ophiobolus graminis in infected wheat straw, whether added directly to the straw or to the surrounding soil. When the infected straws were buried in washed quartz sand, 0.5 g. nitrogen per 100 g. air-dry straw was the optimum dressing for longevity of Ophiobolus. Addition of sodium phosphate did not significantly increase longevity.
Nitrogen is considered to prolong the life of Ophiobolus by enabling the mycelium to form new branch hyphae, which can explore unexhausted parts of the substrate; it is suggested that aged mycelium dies from carbohydrate starvation, through exhaustion of the zones of enzymic erosion around the hyphae. This hypothesis is supported by the extended life of the fungus in infected straws that were shaken twice weekly in 3 % dextrose solution.
Ophiobolus was found to survive longer in infected straws buried in a fallow soil than in the same soil under oats, mustard or trefoil; this finding suggests the use of catch crops as competitors with Ophiobolus for soil nitrogen.