Long-term performance of fungicides applied to control eyespot in winter wheat

Fungicide treatments, repeated twice yearly at timings suitable for controlling eyespot, to plots of continuous winter wheat over 17 yr (except for a year in “set‐aside”), resulted in altered population structures of the eyespot pathogens (Tapesia spp.) and performance of the fungicides. Carbendazim failed to control eyespot after 3 yr of the experiment because of rapid selection for resistance. Carbendazim often increased disease compared with that in untreated plots. This may have resulted from control of unknown antagonistic fungi or, possibly, from selection for increased fitness in carbendazim ‐resistant strains. In the final years of the experiment, prochloraz, applied with or without carbendazim, became ineffective. This gradual loss of efficacy is explained by selection in favour of T. acuformis rather than T. yallundae (which had a naturally narrower range of sensitivity levels in its populations) and for strains with least sensitivity within the T. acuformis population, although no resistance to prochloraz was found. Effects of treatments on yield were usually not statistically significant. This may be explained partly by the predominance of the more slowly developing T. acuformis in many of the plots.     

Surveys of diseases of winter wheat in England and Wales, 1976–1988

Samples from 300 – 400 randomly selected winter wheat crops were taken annually at growth stage 73 – 75 from 1976 to 1988 with the exception of 1983 and 1984. The number of samples from each region was proportional to the area of wheat grown in each region. The percentage of the area of the top two leaves affected by diseases, the severity of ear and stem base diseases and, in 6 years, the severity of take-all were recorded. Septoria tritici and Septoria nodorum were, on average, the most severe of the foliar diseases and eyespot (Pseudocercosporella herpotrichoides) was the most severe of the stem base diseases. Regional differences in levels of S. tritici, brown rust, sharp eyespot and nodal fusarium were significant. Cultivar resistance affected disease severity, and previous cropping patterns particularly affected take-all and eyespot. Eyespot and sharp eyespot were less severe in late- than in early-sown crops. The percentage of crops treated with a fungicidal spray increased from 14% in 1976 to over 90% between 1983 and 1985. Use of benzimidazole fungicides applied at growth stage 31 declined, while use of morpholines from flag leaf emergence onwards increased between 1985 and 1988.     

Surveys of diseases of winter barley in England and Wales, 1981–1991

Samples from 200–400 randomly selected winter barley crops were taken annually at growth stage 71–73 from 1981 to 1991, with the exception of 1984 and 1985. The number of samples from each region was proportional to the area of barley growth in each region. The percentage of the area of the top two leaves affected by diseases and the severity of stem base diseases were recorded. Mildew (Erysiphe graminis) was the most widespread of the foliar diseases and in three years (1982, 1986 and 1991) was also the most severe. Rhynchosporium (Rhynchosporium secalis), net blotch (Pyrenophora teres) and brown rust (Puc-cinia hordei) were also prevalent in some years. Of the stem base diseases, fusarium was often the most widespread. Eyespot (Pseudocercosporella her-potrichoides) severity varied widely from year to year ranging from 1.2% of stems affected by moderate or severe symptoms in 1982 to 24.1% in 1988. There were regional differences in the severity of mildew, rhynchosporium, brown rust, halo spot (Selenophoma donacis) and eyespot. Cultivar resistance affected disease severity and previous cropping affected eyespot and less frequently mildew, rhynchosporium and net blotch. Eyespot, and to a lesser extent, sharp eyespot, were less severe in late- than in early-sown crops. The percentage of crops treated with a fungicidal spray increased from 72% in 1981 to 95% in 1991. The use of benzimidazole fungicides for the control of eyespot declined in response to the development of resistance, and more recently the use of prochlo-raz also declined. Broad spectrum DMI fungicides were widely used, and the use of morpholines to improve mildew control increased significantly. The proportion of crops grown from seed treated with a non-mercurial fungicidal seed dressing reached a peak of 47% in 1986 but subsequently declined to 22% in 1990 and 1991.     

Sources of resistance and susceptibility to Septoria tritici blotch of wheat

An association genetics analysis was conducted to investigate the genetics of resistance to Septoria tritici blotch, caused by the fungus Zymoseptoria tritici (alternatively Mycosphaerella graminicola), in cultivars and breeding lines of wheat (Triticum aestivum) used in the UK between 1860 and 2000. The population was tested with Diversity Array Technology (DArT) and simple‐sequence repeat (SSR or microsatellite) markers. The lines formed a single population with no evidence for subdivision, because there were several common ancestors of large parts of the pedigree. Quantitative trait loci (QTLs) controlling Septoria resistance were postulated on 11 chromosomes, but 38% of variation was not explained by the identified QTLs. Calculation of best linear unbiased predictions (BLUPs) identified lineages of spring and winter wheat carrying different alleles for resistance and susceptibility. Abundant variation in Septoria resistance may be exploited by crossing well‐adapted cultivars in different lineages to achieve transgressive segregation and thus breed for potentially durable quantitative resistance, whereas phenotypic selection for polygenic quantitative resistance should be effective in breeding cultivars with increased resistance. The most potent allele reducing susceptibility to Septoria, on chromosome arm 6AL, was associated with reduced leaf size. Genes which increase susceptibility to Septoria may have been introduced inadvertently into UK wheat breeding programmes from cultivars used to increase yield, rust resistance and eyespot resistance between the 1950s and 1980s. This indicates the need to consider trade‐offs in plant breeding when numerous traits are important and to be cautious about the use of non‐adapted germplasm.     

Effects of cultivar and fungicides on stem-base pathogens, determined by quantitative PCR, and on diseases and yield of wheat

The polymerase chain reaction (PCR) was used to identify and quantify all fungal pathogens of wheat (Triticum aestivum) stem bases in nine field experiments at three locations in England. The main aim was to apply quantitative PCR to provide robust data on the efficacy of new fungicides against the individual components of the stem‐base disease complex. Cyprodinil most effectively controlled eyespot by decreasing both pathogens, Tapesia yallundae and T. acuformis (the most widespread species), and sometimes contributed to increased yields. Prochloraz controlled eyespot less consistently, its effectiveness dependent mainly on the presence of T. yallundae or on rainfall events soon after application. Azoxystrobin contributed to yield increases most consistently. Although it decreased sharp eyespot and its pathogen, Rhizoctonia cerealis, these effects were insufficient to account for much of the yield increases. The effects of fungicides on eyespot were sometimes greatest on the most susceptible cultivars. Amounts of Tapesia DNA were usually consistent with cultivar susceptibility ratings. The only pathogens of brown foot rot present in significant amounts were Microdochium nivale vars nivale and majus. They appeared not to affect yield or to respond greatly to fungicides. The susceptibility of cultivars to these pathogens was sometimes similar to their susceptibility to eyespot, suggesting that they may respond to the same host resistance genes or may in some cases be secondary colonisers of eyespot‐infected plants.     

Mapping of QTL conferring resistance to sharp eyespot (Rhizoctonia cerealis) in bread wheat at the adult plant growth stage

Key message

Seven sharp eyespot resistance QTL were detected consistently across five environments and delimited to seven DNA marker intervals, respectively, six of which were independent of plant height and heading time.

Abstract

Sharp eyespot, caused mainly by the soil-borne fungus Rhizoctonia cerealis, is one of the important diseases of bread wheat (Triticum aestivum L.). This disease has escalated into a major threat to wheat production in some regions of the world. Wheat resistance to sharp eyespot can be a potential means to reduce the needs for application of fungicides and agricultural inputs. In the present study, the winter wheat lines, Luke and AQ24788-83, both of which possess quantitative resistance to sharp eyespot, were crossed and a population consisting 241 recombinant-inbred lines (RILs) was constructed. These RILs were assessed for sharp eyespot resistance by conducting five field and greenhouse trials during the period from 2008 to 2012, and they were genotyped with 549 simple-sequence repeat DNA markers. Seven quantitative trait loci (QTL) were detected consistently across the five trial environments to be associated with the sharp eyespot resistance. They were mapped on chromosomes 1A, 2B, 3B, 4A, 5D, 6B, and 7B. Four of these QTL are unequivocally novel, while it is possible that the other three might also be novel. Plant height and heading date of the 241 RILs were recorded in the four field trials. All of the seven disease resistance QTL were independent of plant height and heading time except one that was significantly associated with plant heading time. This association might be attributed genetically to a single QTL, or to different but closely linked QTL. In the case of single QTL, pleiotropism might be involved or the sharp eyespot resistance might be conferred in a physical instead of physiological nature.     

A major QTL for resistance against Fusarium head blight in European winter wheat

We report on the verification of a resistance quantitative trait locus (QTL) on chromosome 1BL (now designated Qfhs.lfl-1BL) which had been previously identified in the winter wheat cultivar Cansas. For a more precise estimation of the QTL effect and its influence on plant height and heading date lines with a more homogeneous genetic background were created and evaluated in four environments after spray inoculation with Fusarium culmorum. Qfhs.lfl-1BL reduced FHB severity by 42% relative to lines without the resistance allele. This QTL did not influence plant height, but significantly delayed heading date by one day. All of the most resistant genotypes of the verification population carried this major QTL displaying its importance for disease resistance. This resistance QTL has not only been found in the cultivar Cansas, but also in the three European winter wheat cultivars Biscay, History and Pirat. A subsequent meta-analysis confirmed the presence of a single QTL on the long arm of chromosome 1B originating from the four mentioned cultivars. Altogether, the results of the present study indicate that Qfhs.lfl-1BL is an important component of FHB resistance in European winter wheat and support the view that this QTL would be effective and valuable in backcross breeding programmes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Identification of QTLs for resistance to Fusarium head blight, DON accumulation and associated traits in the winter wheat variety Arina

Fusarium head blight (FHB) of wheat has become a serious threat to wheat crops in numerous countries. In addition to loss of yield and quality, this disease is of primary importance because of the contamination of grain with mycotoxins such as deoxynivalenol (DON). The Swiss winter cultivar Arina possesses significant resistance to FHB. The objective of this study was to map quantitative trait loci (QTL) for resistance to FHB, DON accumulation and associated traits in grain in a double haploid (DH) population from a cross between Arina and the FHB susceptible UK variety Riband. FHB resistance was assessed in five trials across different years and locations. Ten QTL for resistance to FHB or associated traits were detected across the trials, with QTL derived from both parents. Very few of the QTL detected in this study were coincident with those reported by authors of two other studies of FHB resistance in Arina. It is concluded that the FHB resistance of Arina, like that of the other European winter wheat varieties studied to date, is conferred by several genes of moderate effect making it difficult to exploit in marker-assisted selection breeding programmes. The most significant and stable QTL for FHB resistance was on chromosome 4D and co-localised with the Rht–D1 locus for height. This association appears to be due to linkage of deleterious genes to the Rht-D1b (Rht2) semi-dwarfing allele rather than differences in height per se. This association may compromise efforts to enhance FHB resistance in breeding programmes using germplasm containing this allele.     

Combined application of Bacillus subtilis NJ-18 with fungicides for control of sharp eyespot of wheat

Antagonistic bacterium, Bacillus subtilis NJ-18, can biologically control several plant diseases. A combined strategy of wheat sharp eyespot control consisting of B. subtilis NJ-18 and fungicides was evaluated. Greenhouse and field tests were conducted to determine the effect of combining the B. subtilis NJ-18 with the fungicides flutolanil and difenoconazole for the control of wheat sharp eyespot caused by Rhizoctonia cerealis Van der Hoeven. The sensitivity of B. subtilis NJ-18 to fungicides flutolanil and difenoconazole was tested. The growth of NJ-18 was unaffected by flutolanil in a broth medium, and the survival of NJ-18 spores on wheat seed was unaffected by difenoconazole. In greenhouse experiments, disease control obtained with a combination of NJ-18 and either fungicide was better than the control obtained with the bacterium or fungicides alone, and some combinations of bacterium plus fungicide demonstrated a small synergistic effect in reducing disease. Similar results were obtained with repeated field experiments. In addition to providing a highly effective disease control, the use of combinations enabled a reduced level of fungicide application. Overall, the results suggest that the combination of B. subtilis strain NJ-18 and flutolanil or difenoconazole represents a promising new tool for the control of sharp eyespot of wheat.     

Risk assessment studies on succinate dehydrogenase inhibitors, the new weapons in the battle to control Septoria leaf blotch in wheat

Chemical control of Septoria leaf blotch, caused by Mycosphaerella graminicola, is essential to ensure wheat yield and food security in most European countries. Mycosphaerella graminicola has developed resistance to several classes of fungicide and, with the efficacy of azoles gradually declining over time, new modes of action and/or improvements in host varietal resistance are urgently needed to ensure future sustainable disease control. Several new‐generation carboxamide fungicides with broad‐spectrum activity have recently been introduced into the cereal market. Carboxamides inhibit succinate dehydrogenase (Sdh) of the mitochondrial respiratory chain (complex II) but, because of their single‐site specificity, these fungicides may be prone to resistance development. The objective of this study was to assess the risk of resistance development to different Sdh inhibitor (SDHI) fungicides in M. graminicola. UV mutagenesis was conducted to obtain a library of carboxin‐resistant mutants. A range of SDHI resistance‐conferring mutations was found in Sdh subunits B, C and D. Pathogenicity studies with a range of Sdh variants did not detect any fitness costs associated with these mutations. Most of the amino acid residues identified (e.g. B‐S221P/T, B‐H267F/L/N/Y, B‐I269V and D‐D129E/G/T) are directly involved in forming the cavity in which SDHI fungicides bind. Docking studies of SDHI fungicides in structural models of wild‐type and mutated Sdh complexes also indicated which residues were important for the binding of different SDHI fungicides and showed a different binding for fluopyram. The predictive power of the model was also shown. Further diagnostic development, enabling the detection of resistant alleles at low frequencies, and cross‐resistance studies will aid the implementation of anti‐resistance strategies to prolong the cost‐effectiveness and lifetime of SDHI fungicides.     

Identification of a QTL conferring seedling and adult plant resistance to eyespot on chromosome 5A of Cappelle Desprez

Eyespot is an economically important fungal disease of wheat and other cereals caused by two fungal species: Oculimacula yallundae and Oculimacula acuformis. However, only two eyespot resistance genes have been characterised and molecular markers made available to plant breeders. These resistances are Pch1, introduced into wheat from the relative Aegilops ventricosa, and Pch2, originally identified in the cultivar Cappelle Desprez (CD). There are drawbacks associated with both resistances; Pch1 is linked to deleterious traits carried on the Ae. ventricosa introgression and Pch2 has been shown to have limited effectiveness. An additional resistance has been reported on chromosome 5A of CD that confers resistance to eyespot in adult plants. In the present study, we demonstrate that resistance on this chromosome is effective against both O. yallundae and O. acuformis eyespot pathogens and confers resistance at both seedling and adult plant stages. This resistance was mapped in both seedling bioassays and field trials in a 5A recombinant population derived from a cross between CD and a CD single chromosome substitution line carrying 5A from the susceptible line Bezostaya. The resistance was also mapped using seedling bioassays in a 5A recombinant population derived from a cross between the susceptible line Chinese Spring (CS) and a single chromosome substitution line carrying 5A from CD. A single major QTL on the long arm of chromosome 5A was detected in all experiments. Furthermore, the SSR marker Xgwm639 was found to be closely associated with the resistance and could be used for marker-assisted selection of the eyespot resistance by plant breeders.     

The Development of Eyespot on Seedling Leaf Sheaths in Winter Wheat and Winter Barley Crops Inoculated with W-type or R-type Isolates of Pseudocercosporella herpotrichoides

In 1986–88 the development of eyespot lesions in winter wheat or winter barley differed plots inoculated with W-type isolates of Pseudocercosporella herpotrichoides and plots inoculated with R-type isolates. In the spring of 1986, after a cold winter, the incidence (%shoots infected) and severity (number of leaf sheaths penetrated) of eyespot lesions in wheat before GS 30/31 were greater in plots inoculated with R-type isolates than in those inoculated with W-type isolates. In 1987, after amild winter, eyespot incidence and severity in both wheat and barley were initially greater in W-type plots than in R-type plots. However, by GS 30/31 or 1987. In 1988, when the crop was October-sown, eyespot incidence and severity were greater in W-type than in R-type plots at GS 30/31. Differences in eyespot incidence and severity between W-type and R-type plots were smaller in barley than in wheat. Both the incidence and severity of eyespot were greater in early-sown than in late-sown plots. Seed rate, had little effect on the rate of lesion development in 1987, but in 1988 the rate of penetration was less at the low seed rate for both wheat and barley.     

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.     

Resistance to eyespot of wheat, caused by Tapesia yallundae, derived from Thinopyrum intermedium homoeologous group 4 chromosome

Thinopyrum intermedium was identified previously as resistant to Tapesia yallundae, cause of eyespot of wheat. Using GUS-transformed isolates of T. yallundae as inoculum, we determined that wheat lines carrying Th. intermedium chromosome 4Ai#2 or the short arm of chromosome 4Ai#2 were as resistant to the pathogen as the eyespot-resistant wheat- Th. ponticum chromosome substitution line SS767 (PI 611939) and winter wheat cultivar Madsen, which carries gene Pch1 for eyespot resistance. Chromosome 4E from Th. elongatum and chromosome 4J from Th. bessarabicum did not confer resistance to T. yallundae. Genome-specific PCR primers confirmed the presence of Thinopyrum chromatin in these wheat- Thinopyrum lines. Genomic in situ hybridization using an St genomic probe from Pseudoroegneria strigosa demonstrated that chromosome 4Ai#2 belongs to the J^s genome of Thinopyrum. The eyespot resistance in the wheat- Th. intermedium lines is thus controlled by the short arm of this J^s chromosome. This is the first report of resistance to T. yallundae controlled by a J^s genome chromosome of Th. intermedium.     

Genetic analysis of durable leaf rust resistance in winter wheat

Quantitative resistance that delays the epidemic development of leaf rust in wheat is an important source for durable resistance breeding. The Swiss winter wheat variety ’Forno’ shows a high level of quantitative resistance against leaf rust. This resistance has been effective for more than 10 years and can therefore be considered to be durable. In order to map quantitative trait loci (QTL) for durable leaf rust resistance we analysed 204 F₅ recombinant inbred lines (RILs) of the cross between the winter wheat ’Forno’ and the winter spelt ’Oberkulmer’ for their level of leaf rust resistance (LR) and leaf tip necrosis (LTN) in four different environments. Both traits showed a continuous distribution and were significantly correlated (r=−0.5). Across environments we detected 8 QTL for leaf rust resistance (6 inherited from ’Forno’) and 10 QTL for the quantitative expression of LTN (6 inherited from ’Forno’). Of the 6 QTL responsible for the durable leaf rust resistance of ’Forno’, 1 major QTL coincided with a thaumatin locus on 7BL explaining 35% of the phenotypic variance. Four QTL for LR coincided with QTL for LTN. At these loci the alleles of ’Forno’ increased the level of resistance as well as the extent of LTN, indicating pleiotropy. Received: 1 July 1999 / Accepted: 30 July 1999     

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.     

Semiquantitative,dünnschichtchromatographische Bestimmung von DNOC- und DNBP-Rückständen in Pflanze und Boden

It has been represented the data on the pathogens’ population of eye spot of wheat (Pseudocercosporella spp.) on winter wheat in the Ukraine. They are characterised with different properties of pathogenicity and cultural‐morphological peculiarities. It was marked the interdependence between the growth rate of strains and the properties of pathogenicity in vitro. It was stated the differentiation of the regional varieties of the winter wheat according to their resistance to the disease in different soil‐climatic zones in Ukraine.     

Effect of previous crops on the incidence of eyespot on winter wheat

Surveys of winter wheat from 1939 to 1946 show that eyespot (Cercosporella herpotrichoides Fron.) occurs throughout Britain and that its incidence depends largely on previous cropping and on weather. Examination of 551 crops on land whose cropping for the previous 4 years was known showed that the incidence rose steadily with increasing numbers of preceding wheat and barley crops: where neither crop had been taken for 4 years the proportion of crops with more than 70% infected straws was 2%, rising to 45% where three or four such crops had been taken and the average straws infected rose from 6 to 55%.
The percentage infection to be expected in various groups of crops was calculated from previous cropping; it was compared with the actual infection and so used to assess the importance of other factors in determining the incidence of eyespot. High spring rainfall, early sowing and a dense plant increased incidence and low spring rainfall, late sowing and a thin plant reduced it.
Eyespot was not usually severe on newly ploughed grassland until the third or fourth crop of wheat, but under very wet conditions it was sometimes severe in the second crop.
Oats is much less susceptible than wheat or barley, but some crops were found with a third of their straws infected.
A brief survey of winter wheat in Holland suggested possible causes for the rise and fall of eyespot in recent years and for its present lower incidence there as compared with East Anglia.     

Results of long-term field studies into the effect of strobilurin containing fungicides on the production of mycotoxins in several winter wheat varieties

In total, 23 field trials have been performed during the past 5 years to assess the effect of strobilurin containing fungicides on mycotoxin production in winter wheat. In 85% of the plots, the treatment with strobilurins at growth stages before blossom increased the content of deoxynivalenol in comparison with untreated controls. This effect was intensified by treatments at BBCH stage 45–49 and could be demonstrated in a great number of various cultivars. Application of strobilurin containing fungicides at early growth stages seems to be a risk factor for mycotoxin formation in wheat, especially under favourable conditions forFusarium infections. Presented at the 26^th Mykotoxin-Workshop in Herrsching. Germany, May 17–19, 2004