Tolerance of septoria leaf blotch in winter wheat

For individual varieties, tolerance of septoria leaf blotch was quantified by the slope of the relationship between disease and yield. Variation in disease severity and the associated yield responses were provided across two sites and three seasons of field experiments. Slopes were fitted by residual maximum likelihood for two contrasting models: (i) a fixed-effects model, where no prior assumptions were made about the form of the variety slopes; and (ii) a random-effects model, where deviations in individual variety slopes away from the mean variety slope formed a normal random population with unknown variance. The analyses gave broadly similar results, but with some significant differences. The random model was considered more reliable for predicting variety performance. The effects of disease were quantified as symptom area and green canopy duration. Models of the relationship between symptom area and yield were site-specific. When site effects were not taken into account, these models had poor predictive precision. Models based on the canopy green area gave robust predictions of yield and were not site-specific. Differences in disease tolerance were detected in a comparison of 25 commercial winter wheat varieties. Tolerance was not detected directly through symptom measurements, but instead through measurements of canopy green area, which provides a measurement of the effects of disease that accounts for differences in canopy size across sites and seasons. The varieties showing greatest tolerance tended to have lower attainable yield than the intolerant varieties. Presence of the 1BL/1RS chromosome translocation, which has been reported to increase radiation use efficiency, appeared to be associated with intolerance.     

Position of inoculum in the canopy affects the risk of septoria tritici blotch epidemics in winter wheat

The spread of septoria leaf blotch in wheat caused by Mycosphaerella graminicola is widely reported to depend on the occurrence of splashy rainfall. Previous studies have also implicated an important effect of canopy architecture on the risk of disease spread. This is because architecture affects the proximity of the yield-forming leaves to inoculum present on older diseased leaves within the crop. This study demonstrated that infection of the final three leaves of winter wheat could occur in the absence of splashy rainfall. For cvs Riband and Longbow, the final two leaf layers emerged at a position ≈ 8 cm below older leaves containing sporulating lesions. Under these conditions, infection of new leaves occurred in treatments that simulated dew and nonsplashy rainfall. These treatments resulted in disease incidences of 10–40% above the untreated control on the final two leaf layers. Within a season, the distance between diseased and healthy leaves during the period of stem extension varied substantially across a range of 30 cultivars. While the magnitude of these differences was not the same across seasons, the relative differences between cultivars were generally consistent, suggesting a strong genotype influence on lesion proximity. This study shows how knowledge of the distribution of lesion proximity within a crop can be used to estimate the risk of inoculum transfer for a given maximum splash height. A rapid crop-monitoring method for estimating the distribution of lesion proximity was developed and tested. Lesion proximity was manipulated by plant growth regulator (PGR) treatments. Significant increases in disease, between 14 and 62%, were observed on the upper canopy leaves of plants treated with PGR. The largest differences were observed in treatments where lesion proximity was most affected.     

Sources of resistance to septoria tritici blotch and implications for wheat breeding

Twenty-four wheat cultivars and breeding lines were screened for isolate-specific resistance to septoria tritici blotch (STB) caused by 12 isolates of Mycosphaerella graminicola. New isolate-specific resistances that could be used in wheat breeding were identified. Major sources of resistance to STB used in world breeding programmes for decades, such as Kavkaz-K4500, Veranopolis, Catbird and TE9111, have several isolate-specific resistances. This suggests that 'pyramiding' several resistance genes in one cultivar may be an effective and durable strategy for breeding for resistance to STB in wheat. Several cultivars, including Arina, Milan and Senat, had high levels of partial resistance to most isolates tested as well as isolate-specific resistances. Resistance to isolate IPO323 was common, present in all but one of the major sources of resistance tested. This suggests that resistance to IPO323 may be an indicator of varietal resistance to STB in the field.     

Contributions of disease resistance and escape to the control of septoria tritici blotch of wheat

The contributions of disease escape and disease resistance to the responses of wheat to septoria tritici leaf blotch (STB) were analysed in a set of 226 lines, including modern cultivars, breeding lines and their progenitors dating back to the origin of scientific wheat breeding. Field trials were located in the important wheat-growing region of eastern England and were subject to natural infection by Mycosphaerella graminicola . STB scores were related to disease-escape traits, notably height, leaf spacing, leaf morphology and heading date, and to the presence of known Stb resistance genes and isolate-specific resistances. The Stb6 resistance gene was associated with a reduction of 19% in the level of STB in the complete set of 226 lines and with a 33% reduction in a subset of 139 lines of semidwarf stature. Greater plant height was strongly associated with reduced STB in the full set of lines, but only weakly in the semidwarf lines. Shorter leaf length was also associated with reduced STB, but, in contrast to earlier reports, lines with more prostrate leaves had more STB on average, probably because they tended to have longer leaves. Several lines, notably cvs Pastiche and Exsept, had low mean levels of STB which could not be explained by either escape traits or specific resistance genes, implying that they have unknown genes for partial resistance to STB.     

Early stages of septoria tritici blotch epidemics of winter wheat: build‐up,overseasoning, and release of primary inoculum

Septoria tritici blotch (STB), caused by Mycosphaerella graminicola, is the most prevalent disease of wheat worldwide. Primary inoculum and the early stages of STB epidemics are still not fully understood and deserve attention for improving management strategies. The inoculum build‐up and overseasoning involves various fungal structures (ascospores, pycnidiospores, mycelium) and plant material (wheat seeds, stubble and debris; wheat volunteers; other grasses). Their respective importance is assessed in this review. Among the mechanisms involved in the early stages of epidemics and in the year‐to‐year disease transmission, infection by ascospores wind‐dispersed from either distant or local infected wheat debris is the most significant. Nevertheless, infection by pycnidiospores splash‐dispersed either from neighbouring wheat debris or from senescent basal leaves has also been inferred from indirect evidence. Mycosphaerella graminicola has rarely been isolated from seeds so that infected seed, although suspected as a source of primary inoculum for a long time, is considered as an epidemiologically anecdotal source. Mycosphaerella graminicola can infect a few grasses other than wheat but the function of these grasses as alternative hosts in natural conditions remains unclear. Additionally, wheat volunteers are suspected to be sources of STB inoculum for new crops. This body of evidence is summarized in a spatio‐temporal representation of a STB epidemic aimed at highlighting the nature, sources and release of inoculum in the early stages of the epidemic.     

A comparison between visual estimates and image analysis measurements to determine septoria leaf blotch severity in winter wheat

Methods to estimate disease severity vary in accuracy, reliability, ease of use and cost. Severity of septoria leaf blotch (SLB, caused by Zymoseptoria tritici) was estimated by four raters and by image analysis (assumed actual values) on individual leaves of winter wheat in order to explore accuracy and reliability of estimates, and to ascertain whether there were any general characteristics of error. Specifically, the study determined: (i) the accuracy and reliability of visual assessments of SLB over the full range of severity from 0 to 100%; (ii) whether certain 10% ranges in actual disease severity between 0 and 100% were more prone to estimation error compared with others; and (iii) whether leaf position affected accuracy within those ranges. Lin's concordance correlation analysis of all severities (0–100%) demonstrated that all raters had estimates close to the actual values (agreement: ρ_c = 0·92–0·99). However, agreement between actual SLB severities and estimates by raters was less good when compared over short 10% subdivisions within the 0–100% range (ρ_c = ?0·12 to 0·99). Despite common rater imprecision at estimating low and high SLB severities, individual raters differed considerably in their accuracy over the short 10% subdivisions. There was no effect of leaf position on accuracy or precision of severity estimate on separate leaves (L1–L3). Pursuing efforts in understanding error in disease estimation should aid in improving the accuracy of assessments, making visual estimates of disease severity more useful for research and applied purposes.     

Pyraclostrobin reduces germ tube growth of QoI‐resistant Mycosphaerella graminicola pycnidiospores and the severity of septoria tritici blotch on winter wheat

The effect of the quinone outside inhibitors (QoI) azoxystrobin and pyraclostrobin on yields of winter wheat where QoI resistant Mycosphaerella graminicola isolates were dominant was investigated in field trials in 2006 and 2007. Pyraclostrobin significantly increased yields by 1·57 t ha^?1 in 2006 and 0·89 t ha^?1 in 2007 when compared to the untreated controls, while azoxystrobin only provided a significant increase of 1·28 t ha^?1 in 2006. These yield increases were associated with reduction in septoria tritici blotch (STB) development as determined by weekly disease assessments over a 7 week interval. The effect of pyraclostrobin on STB was studied in controlled environment experiments using wheat seedlings inoculated with individual M. graminicola isolates. Pyraclostrobin significantly reduced STB symptoms by up to 62%, whether applied 48 h pre‐ or post‐ inoculation with resistant M. graminicola isolates containing the cytochrome b mutation G143A. Extremely limited disease (<1%) was observed on similarly treated seedlings inoculated with an intermediately resistant isolate containing the cytochrome b mutation F129L, while no disease was observed on seedlings inoculated with a wild‐type isolate. Germination studies of pycnidiospores of M. graminicola on water agar amended with azoxystrobin or pyraclostrobin showed that neither fungicide inhibited germination of spores of resistant isolates containing the mutation G143A. However, pyraclostrobin significantly reduced germ tube length by up to 46% when compared with the untreated controls. Although the QoIs can no longer be relied upon to provide effective M. graminicola control, this study provides an insight into why QoIs still provide limited STB disease control and yield increases even in situations of high QoI resistance.     

Effect of temperature on latent period of septoria leaf blotch on winter wheat under outdoor conditions

Batches of two winter wheat cultivars (Riband and Apollo) were inoculated with conidia of Mycosphaerella graminicola at weekly intervals over a 2 year period. Following 72 h incubation, plants were placed in ambient temperatures ranging between −7 and 32°C with mean batch temperatures of 2·9–20·2°C. Latent period until the first visible symptoms ranged between 11 and 42 days. The relationship between development of lesions and accumulated thermal time was described using a shifted cumulative gamma distribution model. The model provided good estimates of lesion development with r ² > 0·92 for both cultivars. Base temperatures, below which the pathogen did not develop, were estimated from the model as approximately −2·4°C for the two cultivars. Latent period was estimated as being 250 and 301 degree-days above the estimated base temperature, when defined as time from inoculation to first lesion and time to 50% of maximal lesions, respectively, for cv. Riband. The values for cv. Apollo were similar, but with estimates of thermal time periods c . 5% higher. The relationship between mean temperature and inverse latent period, expressed as days either to first lesion or to 50% of maximal lesions, was best described by a linear regression with r ² > 0·96 for both cultivars. The opportunity for plants to outgrow disease was reduced when prolonged periods of cold temperature occurred, because the base temperature for growth of the pathogen was less than that for the crop.     

Resistance of wheat cultivars and breeding lines to septoria tritici blotch caused by isolates of Mycosphaerella graminicola in field trials

Isolate-specific resistance of 71 cultivars and breeding lines of wheat ( Triticum aestivum ) to septoria tritici blotch was evaluated in six field trials in the Netherlands, Switzerland and the UK between 1995 and 1997. Each plot was inoculated with one of six single-pycnidium isolates of the pathogen Mycosphaerella graminicola . There were strong interactions between wheat lines and M. graminicola and the line-by-isolate interactions were stable over the six trials. Lines with specific resistance or specific susceptibility to each of the isolates were identified. Specific resistance to isolate IPO323 was especially common, being carried by 22 lines from 10 countries. The results confirm that line-by-isolate interactions in septoria tritici blotch of wheat are effective in adult plants in field conditions, and are not simply confined to seedlings. Wheat lines with good, quantitative resistance to all or most isolates were identified, including lines from Brazil, the USA and seven European countries. These may be useful as sources of resistance in wheat breeding.     

Identification and location of Stb9, a gene for resistance to septoria tritici blotch in wheat cultivars Courtot and Tonic

This study reports the discovery of a gene for resistance to septoria tritici blotch (STB) in two spring wheat cultivars, Courtot and Tonic. The gene, named Stb9 , confers resistance to Mycosphaerella graminicola isolate IPO89011. It was mapped by quantitative trait loci (QTL) analysis using an existing map of Courtot × Chinese Spring and was located between markers Xfbb226 (3·6 cM) and XksuF1b (9 cM) on the long arm of chromosome 2B. Markers linked to Stb9 in Courtot were then shown to be linked to resistance to IPO89011 in F₃ families of Tonic × Longbow. Allelism tests in which Tonic was crossed with Courtot confirmed that Tonic has a gene for resistance to IPO89011 at or very close to the Stb9 locus. SSR markers flanking Stb9 may be used in marker-assisted selection to introgress this gene into winter cultivars or in spring wheat breeding programmes outside Europe.     

Identification of isolate-specific and partial resistance to septoria tritici blotch in 238 European wheat cultivars and breeding lines

From a total of 238 European cultivars and breeding lines screened for isolate-specific resistance to septoria tritici blotch (STB) with eight Mycosphaerella graminicola isolates from five different countries, 142 lines were resistant to Ethiopian isolate IPO88004, and 43 lines were specifically resistant to IPO323, with little or no leaf area bearing pycnidia of M. graminicola . These lines probably all have the resistance gene Stb6 . Specific resistances to isolates CA30JI, IPO001, IPO89011, IPO92006 and ISR398 were less common. Seventy-three per cent of the lines were specifically resistant to at least one isolate and 36 lines were resistant to more than one isolate. The line with the greatest number of specific resistances was the spring cultivar Raffles, with five. The most resistant line in which no specific resistance was identified was the Italian landrace Rieti, an ancestor of many modern European wheat cultivars. There was also a wide range of partial resistance among the lines tested, expressed in detached seedling leaves. Information about the resistance of wheat lines to M. graminicola isolates will assist breeders to choose parents of crosses from which progeny with superior resistance to STB may be selected.     

A predictive model for early-warning of Septoria leaf blotch on winter wheat

Disease–weather relationships influencing Septoria leaf blotch (SLB) preceding growth stage (GS) 31 were identified using data from 12 sites in the UK covering 8 years. Based on these relationships, an early-warning predictive model for SLB on winter wheat was formulated to predict the occurrence of a damaging epidemic (defined as disease severity of 5% or > 5% on the top three leaf layers). The final model was based on accumulated rain > 3 mm in the 80-day period preceding GS 31 (roughly from early-February to the end of April) and accumulated minimum temperature with a 0°C base in the 50-day period starting from 120 days preceding GS 31 (approximately January and February). The model was validated on an independent data set on which the prediction accuracy was influenced by cultivar resistance. Over all observations, the model had a true positive proportion of 0.61, a true negative proportion of 0.73, a sensitivity of 0.83, and a specificity of 0.18. True negative proportion increased to 0.85 for resistant cultivars and decreased to 0.50 for susceptible cultivars. Potential fungicide savings are most likely to be made with resistant cultivars, but such benefits would need to be identified with an in-depth evaluation.     

Presence of the Stb6 gene for resistance to septoria tritici blotch (Mycosphaerella graminicola) in cultivars used in wheat-breeding programmes worldwide

Fifty-two wheat cultivars and breeding lines, most of which have been used in breeding programmes worldwide, were tested for isolate-specific resistance to Mycosphaerella graminicola isolate IPO323, which interacts with the Stb6 gene of wheat (first identified in cvs Flame and Hereward) via a gene-for-gene relationship. Twenty-three lines were specifically resistant to this isolate. Sixteen resistant lines were crossed with Flame for a test of allelism. All progeny lines were resistant, suggesting that the 16 parental lines had Stb6 , a gene allelic to it or a gene closely linked to it. In 14 lines, resistance to IPO323 was controlled by Stb6 only. An exception was Kavkaz-K4500 L6.A.4., which has two genes for resistance to IPO323, one of which is Stb6 . The microsatellite marker Xgwm369 was used to examine genetic diversity in the region of the genome containing Stb6 , to which it is closely linked. Eleven alleles of Xgwm369 , with amplified fragments of 10 different sizes, as well as apparent nonamplification of this marker in Bulgaria 88, were detected. Through the use of information about lines' ancestry, combined with Xgwm369 alleles, it was shown that Stb6 entered world wheat-breeding programmes on a minimum of six occasions, and possibly from as many as 11 sources. The presence of Stb6 in both European and Chinese landraces suggests that this gene has been present in cultivated wheat since the earliest times of agriculture.     

Resistance of wheat to septoria tritici blotch (Mycosphaerella graminicola) and associations with plant ideotype and the 1BL−1RS translocation

Septoria tritici blotch (STB) is a major disease of wheat, reaching epidemic proportions in many parts of the world. In several studies, taller, later-maturing cultivars have had lower disease levels. This study was undertaken to investigate the genetic associations of natural field infection by STB with disease-escape mechanisms related to heading date and height components, mainly leaf spacing, in a population where height and maturity are not controlled by major genes. In field trials of a single seed-descent population of a cross between two nonsemi-dwarf cultivars, Apollo (with strong partial resistance to STB) and Thésée (susceptible), conducted over 3 years, there was a negative correlation between STB and heading date. There was no correlation between STB and distance from stem base to leaf 2; and there was an unexpected positive correlation between STB and distance from flag leaf to leaf 2, which contradicted the so-called 'ladder effect' postulated in STB epidemiology. No effect was detected of the presence of the 1BL−1RS translocation on STB levels. The largest single contributor to variation in STB levels was genetic variation between the progeny lines, and the narrow-sense heritability was 42%. These results suggest that breeders can select for STB resistance alongside optimal stature within the range of height which is adaptive in a particular environment.     

Suppression of septoria tritici blotch and leaf rust on wheat seedling leaves by pseudomonads

Application of cells of two isolates of fluorescent pseudomonads from soil to wheat seedlings prior to inoculation with Mycosphaerella graminicola (anamorph, Septoria tritici) or Puccinia recondita f.sp. tritici markedly reduced symptom expression. These Pseudomonas isolates, LEC 1 and LEC 2. also reduced in vitro growth of Geotrichum candidum. Rhizoctonia solani. Sclerotium rolfsii and S. tritici. Growth of the melanin-producing isolate ISR398 of S. tritici was inhibited on silica gel thin-layer chromatograms by compound(s) extracted with diethyl ether from King's Medium B colonized by Pseudomonas isolate LEC 1. The growth of the antagonistic pseudomonads on defined medium was not affected by the following commercial fungicides: benomyl, captafol, chlorothalonil, fenarimol, mancozeb, maneb, metalaxyl, prochloraz, propiconazole, triadimefon, and the herbicides 2,4-D and diclofop-methyl at the recommended concentrations     

Comparative pathogenicity of sexual and asexual spores of Zymoseptoria tritici (septoria tritici blotch) on wheat leaves

Zymoseptoria tritici ascospores and pycnidiospores are considered the main forms of primary and secondary inoculum, respectively, in septoria tritici blotch epidemics. The pathogenicity of the two types of spores of the same genotypic origin were compared through a two‐stage inoculation procedure in controlled conditions. Adult wheat leaves were inoculated with ascospores collected from field sources, yielding 119 lesions; pycnidiospores collected from 12 lesions resulting from these ascospore infections were then used for inoculation. Lesion development was assessed for 5 weeks; latent period, lesion size, and pycnidium density were estimated for different isolates. The latent period was calculated as the maximum likely time elapsed between inoculation and either the appearance of the majority of the sporulating lesions (leaf scale) or the appearance of the first pycnidia (lesion scale). The latent period was significantly longer (c. 60 degree‐days, i.e. 3–4 days) after infection with ascospores than with pycnidiospores. No difference was established for lesion size and density of pycnidia. A comparison with other ascomycete fungi suggested that the difference in latent period might be related to the volume of spores and their ability to cause infection. Fungal growth before the appearance of lesions may be slower after inoculation with an ascospore than with a pycnidiospore. The mean latent period during the very beginning of epidemics, when first lesions are mainly caused by ascospores, may be longer than during spring, when secondary infections are caused by pycnidiospores. Disease models would be improved if these differences were considered.     

Protective effects of a wheat cultivar mixture against splash‐dispersed septoria tritici blotch epidemics

The use of cultivar mixtures to control foliar fungal diseases is well documented for windborne diseases, but remains controversial for splash‐dispersed diseases. To try to improve this strategy, a cultivar mixture was designed consisting of two wheat cultivars with contrasted resistance to Mycosphaerella graminicola , responsible for the rainborne disease septoria tritici blotch (STB), in a 1:3 susceptible:resistant ratio rather than the 1:1 ratio commonly used in previous studies. The impact of natural STB epidemics in this cultivar mixture was studied in field experiments over 4 years. Weekly assessments of the number of sporulating lesions, pycnidial leaf area and green leaf area were carried out on the susceptible cultivar. In years with sufficient STB pressure, disease impacts on the susceptible cultivar in the mixture were always significantly lower than in the pure stand (e.g. 42% reduction of pycnidial leaf area for the three upper leaves in 2008 and 41% in 2009). In years with low STB pressure (2010 and 2011), a reduction of disease impacts was also shown but was not always significant. After major rainfall events, the number of sporulating lesions observed on the susceptible cultivar after one latent period was reduced on average by 45% in the mixture compared to the pure stand. All the measurements showed that a susceptible cultivar was consistently protected, at least moderately, in a mixture under low to moderate STB pressure. Therefore, the results prove that the design of an efficient cultivar mixture can include the control of STB, among other foliar diseases.     

Substantial reductions in winter wheat diseases caused by addition of straw but not manure to soil

Over a period of 3 years, five agronomically distinct crops of winter wheat were grown in plots in which straw (1 kg m⁻²), manure (4 kg m⁻²) or nothing were incorporated into the soil. Plant establishment and height, but not leaf area per tiller, were lower in straw-treated plots. Fertilizer regimes differed between years. Soil and leaf nitrogen were recorded; there was no obvious link between N and any disease or soil amendment. The numbers of leaf layers scorable for disease were similar in all treatments. At the end of the season, plants from straw-treated plots had consistently reduced septoria tritici blotch (caused by Mycosphaerella graminicola ), powdery mildew [caused by Erysiphe ( Blumeria ) graminis ], brown rust (caused by Puccinia recondita ) and foot rot (caused by Fusarium spp.). Early on, M. graminicola was worse in straw-treated plots. In manure-treated plots, P. recondita was reduced but effects on other diseases were inconsistent and slight. A fungicide, chlorothalonil, was applied in one crop; its effects did not interact with those of other treatments. Mycosphaerella graminicola was not suppressed by straw in outdoor pot experiments. Late in the season, straw-treated plants had significantly higher leaf silica ( P  < 0·01). In a glasshouse experiment, plants supplied with silicon had less E. graminis infection ( P  < 0·001) and higher leaf silica, but effects on M. graminicola were inconsistent. A prior inoculation of M. graminicola primed plant defences against a subsequent attack of E. graminis , but only in the presence of adequate Si. It is postulated that straw acts in the field by increasing Si availability.     

Effects of straw and silicon soil amendments on some foliar and stem-base diseases in pot-grown winter wheat

Four foliar and two stem-base pathogens were inoculated onto wheat plants grown in different substrates in pot experiments. Soils from four different UK locations were each treated in three ways: (i) straw incorporated in the field at 10 t ha⁻¹ several months previously; (ii) silicon fertilization at 100 mg L⁻¹ during the experiment; and (iii) no amendments. A sand and vermiculite mix was used with and without silicon amendment. The silicon treatment increased plant silica concentrations in all experiments, but incorporating straw was not associated with raised plant silica concentrations. Blumeria graminis and Puccinia recondita were inoculated by shaking infected plants over the test plants, followed by suitable humid periods. The silicon treatment reduced powdery mildew ( B. graminis ) substantially in sand and vermiculite and in two of the soils, but there were no effects on the slight infection by brown rust ( P. recondita ). Phaeosphaeria nodorum and Mycosphaerella graminicola were inoculated as conidial suspensions. Leaf spot caused by P. nodorum was reduced in silicon-amended sand and vermiculite; soil was not tested. Symptoms of septoria leaf blotch caused by M. graminicola were reduced by silicon amendment in a severely infected sand and vermiculite experiment but not in soil or a slightly infected sand and vermiculite experiment. Oculimacula yallundae (eyespot) and Fusarium culmorum (brown foot rot) were inoculated as agar plugs on the stem base. Severity of O. yallundae was reduced by silicon amendment of two of the soils but not sand and vermiculite; brown foot rot symptoms caused by F. culmorum were unaffected by silicon amendment. The straw treatment reduced severity of powdery mildew but did not detectably affect the other pathogens. Both straw and silicon treatments appeared to increase plant resistance to all diseases only under high disease pressure.