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.     

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.     

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.     

Comparison of natural and artificial epidemics of take-all in sequences of winter wheat crops

Winter wheat was grown for six successive years (Expt 1) and for three successive years (Expt 2) in field experiments on different soil types. Artificial inoculum of the take-all fungus (Gaeumannomyces graminis var. tritici cultured on autoclaved oat grains) was incorporated in the soil of some of the plots just before, or at, sowing of the first winter wheat crop. Expt 1 tested the incorporation of similar amounts of inoculum (212 kg ha^-1) at different depths. Expt 2 tested different amounts of inoculum at the same, shallow depth. Early sowing (September), late sowing (October) and spring inoculation were additional treatments, applied to the first crop only, in Expt 2. Seasonal factors apart, the disease outcome in the first year after inoculation depended on amounts and placement of applied inoculum, as well as date of sowing. Deeper inoculum resulted in less disease (Expt 1). Severe take-all was produced in Expt 2 by incorporating inoculum shallowly in sufficient quantities (400 kg ha^-1 or more). Less inoculum (200 kg ha^-1) generated less disease, especially in earlier-sown plots. Differences in disease amongst inoculum treatments were greatest in the first year and diminished subsequently, particularly where sowing had been early in the first year. In Expt 1, where first crops exposed to artificial inoculum developed moderate-to-severe disease, disease in subsequent second and/or third crops was less. In the fourth crop a second peak of disease occurred, coinciding with a first peak in sequences without added inoculum. Take-all decline (TAD) appeared to be expressed in all sequences thereafter. In Expt 2 in sequences without added inoculum, TAD occurred after a peak of disease in the second crop. Where 400 kg ha^-1 or more of inoculum were added, disease was severe in the first year and decreased progressively in successive years. Disease was less patchy in plots that received artificial inoculum. However, it remains uncertain mat severe disease caused by artificial inoculation achieved an early onset of true TAD. The infectivity of the top 12 cm of soil in the first 3 yr of Expt 1, determined by bioassay, depended on the depth of added inoculum and amount of disease in subsequent crops. However, at the time of the naturally occurring peak of disease severity (in either inoculated or non-inoculated plots) it did not predict either disease or TAD. Differences and similarities amongst epidemics developing naturally and those developing from different amounts and placement of applied inoculum have been revealed. The epidemiological implications of adding inoculum and the potential value of artificially-created epidemics of take-all in field trials are discussed.     

The microbiology of soils under successive wheat crops in relation to take-all disease

Abstract Thirty-eight wheat fields in southern England were sampled in an attempt to correlate the amount of take-all disease with 35 microbiological and chemical measurements of soil. There was little correlation between field take-all and pot tests to determine soil infectivity. Myxogastrids were important components of the soil population, being up to half of the amoebal population, and most soils contained dictyostelids, reticulate amoebae and myxobacteria. Amoebae, ciliates, bacteria and saprophytic fungi were recorded for all soils. pH was a major determinant of soil populations, being clearly correlated with fungal abundance and with numbers of ciliates, dictyostelids and bacteria. Principal component analysis separated dictyostelids from the other soil amoebae and again showed the importance of pH in determining soil microbial populations. Take-all was negatively correlated with soil fertility and positively related to nematodes and myxobacteria, but this was probably an effect of take-all, and represented saprophytic growth on dead roots rather than being a cause. Reticulate amoebae and dictyostelids were both correlated with low levels of take-all. This study emphasises the large number of interrelated populations of soil microorganisms which could have an effect on the severity of take-all infections.     

The potential of non-pathogenic Gaeumannomyces spp., occurring naturally or introduced into wheat crops or preceding crops, for controlling take-all in wheat

Take‐all disease (Gaeumannomyces graminis var. tritici) in wheat crops is known to be suppressed by naturally occurring antagonistic fungi, closely related to the pathogen, that infect grasses and cereals. This form of suppression was re‐investigated because of the changing importance and role of grass weeds and grass covers in arable farming. Natural populations of the competitive fungus Gaeumannomyces cylindrosporus, allowed to develop under rye‐grass, were more effective than artificially introduced populations in suppressing the development of take‐all in following wheat crops. To be effective, the antagonist needs to be present before the start of wheat cropping. Introducing G. cylindrosporus, but not G. graminis var. graminis (a potential antagonist that is faster growing), into a previous crop, or just after the previous crop, sometimes suppressed take‐all, but the effect was small. It is concluded that, for any future attempts at biocontrol by these fungi, they should be introduced into a preceding crop not susceptible to take‐all. Take‐all inoculum in the soil should be at a minimum and effective hosts of the take‐all pathogen must not be present as weeds or volunteers.     

Nitrate production in the rhizosphere of Plantago species

Summary The production of nitrate in an old established dune grassland soil and its uptake by plants was studied by comparing amounts of mineral nitrogen and numbers of nitrifying bacteria in the rhizosphere on the one hand, and on the other accumulated nitrate and levels of nitrate reductase (NaR) of individual plants of three Plantago species,i. e., P. major, P. lanceolata andP. coronopus. For these three Plantago species andP. media basal levels of NaR in the absence of nitrate were determined in plants grown in culture solutions. The basal NaR levels ofP. major andP. media (species occurring on nutrient-rich soils) were significantly higher than those ofP. lanceolata andP. coronopus (species found on nutrient-poor soils). NaR activity increased in the presence of nitrate and was suppressed by ammonium.From the numbers of nitrifying bacteria in the rhizosphere and NaR activity in the leaves it was concluded that nitrate was produced in the root environments of the three Plantago species and that the compound was taken up by the plants. NaR activities and numbers of nitrifying bacteria were higher for individuals ofP. major than for those ofP. lanceolata andP. coronopus. No correlation was found between the ammonium levels and the numbers of nitrifying bacteria in the soil, and no indications of inhibition of nitrifying bacteria in the rhizosphere were obtained. For individuals ofP. lanceolata a correlation was found between the numbers of nitrifying bacteria in the soil and NaR activity in the leaves. The results are discussed in relation to the ecological habitats of the three species.Grassland Species Research Group Publication No.38.     

Observations on take-all and eyespot diseases of wheat in Yorkshire

Wheat crops were surveyed in Yorkshire from 1944 to 1946 on farms where crops were reported to be unsatisfactory. Take-all ( Ophiobolus graminis (Sacc.) Sacc.) and eyespot ( Cercosporella herpotrichoides Fron.) were both found to be present; the latter is the most serious trouble on the better wheat lands.
The variation in the incidence of these diseases on selected farms during the three seasons has been compared.
The effect of rotation has been examined; both diseases were found to be encouraged by too frequent cropping with wheat or barley. The incidence of disease in wheat crops following a 1-year seeds ley was found to be influenced by the nurse crop used to undersow the seeds, and also by the time at which the ley was ploughed up. Oats were found to be preferable to either wheat or barley as a nurse crop, and in 1946 late ploughing considerably reduced the amount of eyespot disease present in the following wheat crop.
A 1-year ley is considered to be of too short a duration to ensure the disappearance from the soil of Ophiobolus graminis and Cercosporella herpotrichoides surviving from the previous crop, and a period of 2–3 years is suggested as being desirable.     

Nitrate leaching stimulates subsurface root growth of wheat and increases rhizosphere alkalisation in a highly acidic soil

Subsurface acidity is a major factor limiting crop yield in some agricultural soils. The surface application of lime has limited effect on the subsurface acidity due to the slow downward movement, while deep incorporation of lime is costly. This paper tested the concept of biologically ameliorating subsurface acidity in a highly acidic soil through the net uptake of anions by plant roots. Nitrogen was supplied to the top soil (0–10 cm) as Ca(NO₃)₂ at rates equivalent to 30–240 kg N ha^?1. Four water levels were imposed (40, 60, 80 and 100% of field capacity). Aluminium-tolerant wheat was grown for 58 days. The high N and high water treatments stimulated root growth below 15 cm, which in turn increased N capture, resulting in a greater excess anion uptake over cations and thus alkalisation of subsurface soil layers. This study suggests that it is feasible to exploit the process of nitrate uptake by an aluminium-tolerant wheat genotype to increase pH in acidic subsoil.     

Nitrate supply induces changes in polyamine content and ethylene production in wheat plants grown with ammonium

In order to explore the mechanisms of nitrate's beneficial effect on ammonium-grown plants, we investigated the effects of nitrate on free and conjugated polyamine plant content and ethylene biosynthesis in wheat (Triticum aestivum L.) plants grown with ammonium nutrition. Two different doses of nitrate (100 microM and 5 mM) were supplied to ammonium-fed plants, in order to determine whether the effects of nitrate require significant doses (nutritional character), or can be promoted by very low doses (pseudo-hormonal character). Our results showed that nitrate's effects on putrescine, spermidine and spermine contents of ammonium-grown plants tended to follow the pattern associated with strict nitrate nutrition. Both low (100 microM) and high (5 mM) nitrate doses caused a rapid and significant increase in free spermidine content in roots and shoots, which was well correlated with reduced root ethylene production. In shoots, this increase in free spermidine was correlated with changes in the conjugation pattern, while in roots these changes appear to be due to alternative mechanisms. On the other hand, no clear relationship between the supply of a lower dose of nitrate (100 microM) and a reduction of free putrescine content was observed. With higher doses of nitrate (5 mM) we observed a reduction of free putrescine content that was well correlated with increases in its conjugated forms. In conclusion, nitrate's effects on putrescine, spermidine and spermine contents of ammonium-fed plants tended to follow the pattern associated with strict nitrate nutrition, corroborating its beneficial effect.     

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.     

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.     

Effects of foliar sprays of azoxystrobin on take-all in wheat

Average percentages of winter wheat plants with severe take‐all were decreased by up to half by azoxystrobin applied as foliar sprays in four field experiments. Decreased take‐all in three of the experiments was associated with increased grain yield but effects on other diseases may have contributed to these responses. Standard fungicide sprays were ineffective. The effects differed, but not consistently, among different cultivars that were tested in three of the experiments. One, two or three sprays of azoxystrobin or kresoxim‐methyl, in autumn, spring or summer, were tested in the fourth experiment. Unlike azoxystrobin, kresoxim‐methyl had no consistent effects but a smaller amount was applied. Two or three sprays of azoxystrobin were more effective than a single spray but their timing was unimportant. Such control of a root disease by a foliar‐applied fungicide is unusual but may help to explain some of the unexpectedly large yield responses to azoxystrobin that have been reported. This relatively broad‐spectrum fungicide may have the potential to contribute to the practical management of take‐all but further research is needed to determine how best to exploit its effects consistently.     

Natural suppression of take-all disease of wheat in Montana soils

This research was initiated to determine whether soils suppressive to take-all of wheat caused by Gaeumannomyces graminis var. tritici (Ggt) occur in Montana, and to identify the organisms most likely involved in this suppression. From an initial screening of eight soils collected from different wheat growing areas of Montana, two were highly suppressive to take-all. Microbial characterization of these soils indicated that different mechanisms were involved in the suppression. In Larslan soil, mycoparasitism appeared to be the main mechanism. Two different fungi with exceptional ability to reduce the severity of take-all were isolated from this soil. One of these fungi could parasitize the hyphae of Ggt. Field tests with these fungi in Ggt infested soil showed increases of over 100% in both harvestble tillers and grain yield as compared to treatments without these two fungi. In tests with 48 different bacteria and 10 actinomycetes from Larslan soil, none were able to consistently reduce severity of take-all alone, or in mixtures. In Toston soil, antibiosis by actinomycetes and perhaps the involvement of Pseudomonas spp. in production of antibiotics and/or siderophores appeared to be the most likely mechanisms involved in take-all suppression. Increases in shoot dry weight over that in the Ggt infested control using mixtures of pseudomonads and actinomycetes ranged from 25% to 87%. Actinomycetes added individually or in mixtures to soil infested with Ggt consistently reduced the severity of the disease to a greater extent than did mixtures of Pseudomonas spp.     

The use of ammonium fertilisers to reduce the severity of take-all (Gaeumannomyces graminis var. tritici) on wheat in Western Australia

In field experiments in Western Australia ammonium fertilisers significantly reduced the severity of root damage to wheat ( Triticum aestivum ) caused by Gaeumannomyces graminis var. tritici. A series of response lines depicting the effect of sources and rates of nitrogen on take-all are presented. In general, the higher the rate of ammonium nitrogen used, the greater the reduction in the severity of take-all. Ammonium sulphate and ammonium sulphate/mono-ammonium phosphate mixture were more effective in reducing disease than di-ammonium phosphate, urea or ammonium nitrate. Sodium nitrate did not reduce take-all.     

小麦远缘杂交后代对小麦全蚀病抗病性研究

研究了小麦—华山新麦草衍生系和小麦—簇毛麦衍生系19份材料苗期对小麦全蚀病菌禾顶囊壳小麦变种的抗病性,对9份抗病性较高的材料进行田间全生育期抗病性测试。结果表明：9份材料在不同生育期抗病性表现不同,苗期抗病性均高,越冬期无死亡;V2代换系、H922—9—12和V9125—2在返青期抗病性较高,病根严重度低于5．5％;H922—9—12和V9129—1在扬花期抗病性表现较好,成穗率可达80％;H922—9—12和V2代换系在成熟期抗病性较高,白穗率在13．8％以下。对照小偃6号高度感病,蒙燕94—4高度抗病。     

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.