Adult plant resistance to powdery mildew (Erysiphe graminis)in three barley cultivars

The mechanism(s) of adult plant resistance (a.p.r.) to Erysiphe graminis f. sp. hordei were examined in two spring barley cultivars, Athos and Porthos, which possess similar identified mildew resistance genes and seedling resistance, but differential field resistance. These cultivars were compared with Golden Promise, a universally susceptible cultivar. Differential a.p.r. to a compatible race was detected as a reduction in the number and sporulation of colonies on later-formed Athos leaves. Golden Promise exhibited some a.p.r., but much less than either Athos or Porthos. Differential a.p.r. was expressed at several infection stages, viz. appressorial formation, penetration and sporulation. Epidermal hypersensitivity occurred infrequently in all leaf position/cultivar interactions. Papillae were associated with >90% of all penetration failures. The incidence of papillae in successful penetrations at 96 h was 20% on later-formed leaves of Athos compared to c. 60% on all other leaves.     

Effects of nitrogen fertilization and cultivar on the damage relation of powdery mildew (Erysiphe graminis) in winter wheat

The damage relation of powdery mildew in winter wheat was studied in field experiments in 1981 and 1983, in the Netherlands. No firm conclusion was obtained on the effect of nitrogen supply (175 and 235 kg ha^–1 N totally) on the damage relation. The relation was not affected by cultivars (four) and did not differ significantly between both years. The measured relation averaged 0.0125 kg are^–1 damage per pustule-day mildew per leaf, at yields of 70–90 kg are^–1. The effect of the fungicide triadimefon on yield could be ascribed to its effect on diseases. The vertical distribution of mildew in the crop was described.Samenvatting In 1981 en 1983 werd in veldproeven met wintertarwe de relatie tussen schade (in kg are^–1) en meeldauw onderzocht. Of de relatie onafhankelijk is van de stikstofbemesting (175 en 235 kg ha^–1 N totaal), kon niet met zekerheid worden vastgesteld. De schaderelatie werd niet significant beïnvloed door de vier getoetste rassen. In de twee jaren werd een vergelijkbare schade van gemiddeld 0.0125 kg are^–1 per puistdag meeldauw per blad gemeten, bij een opbrengstniveau van 70–90 kg are^–1. Tevens werd de locatie van meeldauw in het gewas beschreven.     

Measures of disease intensity in powdery mildew (Erysiphe graminis) of winter wheat. 1. Errors in estimating pustule number

Assessments of pustule number and severity of powdery mildew on winter wheat in the Netherlands were made in commercial fields and in experimental plots. The sample variance (s²) of the number of pustules per leaf (m) was fairly constant over years, varieties, growth stages and leaf postitions, but depended strongly on the average pustule number: . The effect of sample size on the precision of the estimate is discussed and it is concluded that it is difficult to estimate low disease intensities accurately. Estimates are given for the detection level of pustule counts in relation to sample size.Mildew intensity on the lower surface of leaves can be estimated from the intensity on the upper surface. This method reduces the duration of the observation, but introduces an additional error. At low disease intensities and small sample sizes this method is more efficient than sampling mildew on both surfaces of leaves. The common practice of assessments of the upper surface of leaves only may not be the most efficient method.Samenvatting Aantallen puistjes meeldauw per blad werden geteld in praktijkpercelen en veldproeven met wintertarwe. De steekproefvariantie van het aantal puistjes was tamelijk constant in de jaren, rassen, gewasstadia en bladposities, maar was sterk afhankelijk van het gemiddeld aantal puistjes . Het effect van de steekproefgrootte op de nauwkeurigheid van de schatting wordt besproken en het blijkt dat het moeilijk is om lichte aantastingen nauwkeurig te schatten. Er worden schattingen gegeven van de detectiegrens in afhankelijkheid van de steekproefgrootte.Meeldauwaantastingen aan de onderkant van het blad, kunnen worden geschat uit de aantasting op de bovenkant van het blad. Deze methode levert een tijdsbesparing op, maar ook een extra onnauwkeurigheid. Alleen bij lichte aantastingen en kleine steekproeven is deze methode efficiënter dan een directe tweezijdige bemonstering. Het schatten van meeldauw op de bovenkant van bladeren is, hoewel algemeen gebruikelijk, waarschijnlijk niet de meest efficiënte methode.     

Fungicide sensitivity of populations of wheat powdery mildew (Erysiphe graminis f.sp. Tritici) in Central Europe in 1993

In 1993 we observed the sensitivity of wheat powdery mildew populations (Erysiphe graminis DC f.sp. tritici Marchal) from the Czech Republic, Austria, Hungary and Slovakia to the fungicides triadimenol, tebuconazole, propiconazole, flutriafol and fenpropimorph. The highest resistance value was shown to triadimenol, which attained a mean resistance factor (MRF) of 29 (expressing how many times the population is more resistant than are standard sensitive isolates) in the mildew population from the Czech Republic. The mildew populations from eastern Slovakia and eastern Hungary, populations geographically isolated from the other populations, showed very high sensitivity to all fungicides tested. There was most sensitivity to fenpropimorph (smallest MRF values) compared with the other fungicides. Cross-resistance was established among all triazoles used, but not between triazoles and fenpropimorph. Sensitivity of wheat powdery mildew populations from Central Europe to these fungicides is considered adequate, and the development of resistance has shown a decreasing tendency in recent years.     

Threshold values for chemical control of powdery mildew (Erysiphe cruciferarum) on Brussels sprouts

To establish control thresholds for chemical control of powdery mildew (Erysiphe cruciferarum) on Brussels sprouts, mildew intensity on leaves and buds was observed on the cultivars Lunet, Tardis and Asgard during three years in unsprayed plots. Mildew infection on the leaves was observed from late August onwards, increasing to moderate or high levels. In one year light infestation of the buds was observed, but no reduction in quality occurred. These preliminary results indicate, that from late August onwards the following levels of leaf injury by powdery mildew can be tolerated: T=5+0.42^*(Julian date — 235), in which T is the tolerable leaf injury in percentage leaf area covered. When sampling the crop to assess powdery mildew infection, care must be taken that leaves are sampled from all stem positions, as top leaves tend to be much less infected.     

Components of quantitative resistance to powdery mildew (Erysiphe pisi) in pea (Pisum sativum)

Components of quantitative resistance in pea ( Pisum sativum ) to Erysiphe pisi , the pathogen causing powdery mildew, were investigated. Conidium germination, infection efficiency, latent period and conidium production dynamics on cv. Quantum (quantitatively resistant) were compared with those on Pania and Bolero (susceptible). There was an additional comparison in conidium germination experiments with the resistant cv. Resal. Quantitative resistance in Quantum did not affect conidium germination, but infection efficiency of conidia on this cultivar was 34% less than on the susceptible Pania. More conidia germinated on 5-day-old leaflets than on 15-day-old leaflets but the age of the plant did not affect percentage germination or infection efficiency. The length of the latent period did not differ between cultivars. Total conidium production (AUC) per unit leaflet area on Quantum was 25% less than on Pania. The maximum conidium production per day (CMAX) per unit leaflet area on Quantum was 33% less than on Pania. The time to maximum conidium production per day (TMAX) was 10% longer on Quantum than on Pania. The cv. Bolero, reported to be susceptible, also showed some degree of quantitative resistance, but this differed from that of Quantum. Total conidium production was less on Bolero than on Quantum, but the conidia on Bolero were produced sooner, and for a shorter period, than on Quantum. The stability of these responses was tested by analysing components in three different temperature regimes and testing for interactions with temperature, and with leaflet age. Temperature affected all conidium production variables. AUC per leaflet area was nearly seven times as great and CMAX nearly 15 times greater at 23°C than at 13°C. TMAX increased by 1.5 times when temperature increased from 13°C to 18°C or 23°C. Several interactions occurred and these are described.     

The effect of fungal density on fungicide dose–response curves in barley powdery mildew (Erysiphe graminis f.sp. hordei)

The effect of inoculation density on fungicide dose–response curves and the estimated ED₅₀ value (the fungicide concentration needed to halve the infected leaf area relative to the infected area of leaves that had not been treated with a fungicide) were investigated theoretically, and predictions were tested experimentally using powdery mildew of barley ( Erysiphe graminis [syn. Blumeria graminis ] f.sp. hordei ). A host leaf was assumed to consist of a number of independent compartments, in each of which only a single fungal spore could germinate successfully. The number of fungal spores landing in a compartment was assumed to be Poisson-distributed. The spores were assumed to vary in their sensitivity toward a fungicide, and the sensitivity of the spore population was assumed to be normally distributed around a mean sensitivity. Under these assumptions, the ED₅₀ value was shown to be a positively biased estimate of the mean sensitivity in the fungal population, and the bias increased with inoculum density. Consequently, the estimated ED₅₀ value is expected to vary between experiments and laboratories if the inoculation density varies over a considerable range. The correlation between inoculum density and estimated ED₅₀ value is expected to be strongest when the assayed fungal population consists of several genotypes differing in sensitivity. This expectation was tested by estimating the ED₅₀ values for one barley powdery mildew isolate at different inoculation densities for three different fungicides. These ED₅₀ values were positively correlated strongly with the inoculum density for fenpropimorph and more weakly for triadimenol, whereas no correlation was observed for propiconazole.     

Effect of powdery mildew (Erysiphe pisi) on nodulation and nitrogenase activity in pea (Pisum sativum)

The effects of powdery mildew ( Erysiphe pisi) disease of pea (Pisum sativum) on the number and size of root nodules and on nitrogenase activity were investigated in pea plants grown in pots and in the field. There were significant reductions in nodulation and in the size of root nodules as well as in nitrogenase activity in infected plants.     

Virulence phenotypes in powdery mildew (Blumeria graminis) populations and resistance genes in triticale (x Triticosecale)

Triticale (xTriticosecale) is a new host for powdery mildew (Blumeria graminis) being not infected by this pathogen in Germany before 2001. To evaluate population structure of the pathogen and race-specific resistances in the host, 694 isolates were collected in 12 states of Germany in the years 2007 to 2009 on triticale. They were tested by a newly developed initial differential set of 20 triticale cultivars. Corresponding virulences were found for all differentials except for cultivar Grenado. In total, 272 different virulence phenotypes (=pathotypes) were detected. The virulence complexity of the isolates ranged from 6 to 19 of 20 possible virulences with a mean of 15. In all years, a high level of diversity of the powdery mildew populations was observed with Simpson indices in the range 0.95 to 0.97. The distribution of the pathotypes was even across Germany with an Evenness index in the range 0.82 to 0.88. A set of 19 isolates with different virulence pathotypes and 10 cultivars were selected to be used to identify race-specific resistances of triticale cultivars and breeding lines. Some cultivars susceptible to most of the isolates in seedling stage were moderately resistant in adult-plant stage. The high diversity and complexity of the pathotypes found in German powdery mildew populations as well as an increasing acreage of only a few dominating triticale cultivars accelerate the adaptation of the pathogen to race-specific host resistances suggesting restricted durability only. More durable resistance might be achieved by combining new effective race-specific (qualitative) resistance genes with race-nonspecific (quantitative) resistances effective in the adult-plant stage that are already available.     

Cytological studies on the mode of action of systemic fungicides on the host pathogen complex barley-powdery mildew (Erysiphe graminis f. sp. hordei marchal)

The effects of systemic fungicides on the host-pathogen interface between barley and powdery mildew were studied using cytological methods and specific staining procedures. Propiconazole, etaconazole, bitertanol, triforine, imazalil, nuarimol, tridemorph and fenpropimorph, which are all inhibitors of sterol biosynthesis, stopped growth of the fungus 2 days after application and led to deposits of plant origin around haustoria; these contained polysaccharides, especially callose. This encapsulation resembled the reaction often observed in resistant plants. In contrast, pyrazophos and ethirimol, applied at reduced concentrations, inhibited the fungus at the haustorial stage but did not induce encapsulation. These results suggest that encapsulation is not simply a reaction to the previously damaged fungus, but may be an indirect effect of sterol biosynthesis-inhibiting fungicides on the host metabolism which is elicited by the fungus. The results are discussed in the context of previous observations on the modes of action of these fungicides.     

Spatial and temporal spread of powdery mildew (Erysiphe pisi) in peas (Pisum sativum) varying in quantitative resistance

A field experiment was conducted to assess the progress in time and spread in space of powdery mildew (caused by Erysiphe pisi ) in pea ( Pisum sativum ) cultivars differing in resistance to the disease. Disease severity (proportion of leaf area infected) was measured in 19 × 23 m plots of cultivars Pania and Bolero (both susceptible) and Quantum (quantitatively resistant). Inoculum on infected plants was introduced into the centre of each plot. Leaves (nodes) were divided into three groups within the canopy (lower, middle, upper) at each assessment because of the large range in disease severity vertically within the plants. Disease severity on leaves at upper nodes was less than 4% until the final assessment 35 days after inoculation. Exponential disease progress curves were fitted to disease severity data from leaves at middle nodes. The mean disease relative growth rate was greater on Quantum than on Pania or Bolero, but it was delayed, resulting in an overall lower disease severity on Quantum. Gompertz growth curves were fitted to disease progress on leaves at lower nodes. Disease progress on Quantum was delayed compared with Pania and Bolero. The average daily rates of increase in disease severity from Gompertz curves did not differ between the cultivars on these leaves. Disease gradients in the plots from the inoculum focus to 12 m were detected at early stages of the epidemic, but the effects of background inoculum inputs and the rate of disease progress meant that these gradients decreased with time as the disease epidemic intensified. Spread was rapid, and there were no statistically significant differences between cultivar isopathic rates (Pania 2.2, Quantum 2.9 and Bolero 4.0 m d⁻¹).     

Virulence spectrum in populations of the barley powdery mildew pathogen, Erysiphe graminis f. sp. hordei in Tunisia and Morocco in 1992

The virulence of 57 single colonies of Erysiphe graminis f. sp. hordei from Tunisia and Morocco was investigated using the Pallas near-isogenic differential set. Virulence patterns in general were similar in both countries. Va8, Va10+Du2, V41/145, VLa and VRu2 appear to be common in the region. The resistance alleles Mla7 and Mla9, in combination with other resistances, were highly effective against the isolates tested. No virulence against mlo and Mla9 + k was detected in Tunisia. No virulence on Mla7 and Mla9 was detected in Morocco. The frequency of virulence against several resistances was significantly higher in Morocco than in Tunisia. On the other hand, virulence against other resistances was higher in Tunisia than in Morocco. Three isolates from Morocco infected mlo to an extent greater than previously described. All pathotypes were unique. An attempt is made to interpret the results by comparison with pathotype evolution in Europe.     

Measuring the sterol content of barley leaves infected with powdery mildew as a means of assessing partial resistance to Erysiphe graminis f.sp. hordei

Barley leaf segments of several partially resistant cultivars were inoculated with different strains and combinations of strains of Erysiphe graminis. Infection level was assessed by counting colony number, by measuring sporulating area using an image analyser, and by extracting and quantifying sterols. Good correlations were obtained between data from all three methods. One cultivar contained less sterol than expected from the number of colonies present but some of the resistance in that cultivar was known to be due to reduced colony size. Image analysis was unable to discern the reduced colony size component of resistance. Scanning electron microscopy revealed that single colonies may originate from more than one conidial infection. Sterol analysis was the most representative method of comparing the reaction of diverse host genotypes to mildew infection.     

Resistance to powdery mildew (Blumeria graminis f.sp. avenae) in oat seedlings and adult plants

In this work, 165 Avena sativa and Avena byzantina accessions were screened for resistance to powdery mildew caused by Blumeria graminis f.sp. avenae and the defence mechanisms of resistant plants were further characterized. Ten resistant and moderately resistant accessions were selected according to macroscopic assessment. A detailed histological study of selected genotypes showed a range of defence mechanisms, acting alone or in combination, that impeded fungal development at different stages. Since the resistance observed in the collection was scarce, a study of adult plant resistance was carried out in 45 genotypes selected from field trials. Nine oat landraces and two commercial varieties showed very high levels of adult plant resistance. A detailed study of the components of the adult plant resistance revealed a high increase of penetration and post‐haustorial resistance in the fifth compared to the first leaves. Identification of the resistance sources and characterization of underlying defence mechanisms will be useful for future breeding programmes and for further cellular and molecular studies to unravel the genetic basis of resistance, in this species in particular and in cereal–powdery mildew interactions in general.     

Inhibitory effects of essential oil components extracted from celery (Apium graveolens) on the growth of Septoria apiicola, causal agent of leaf spot disease

Leaf spot disease of celery ( Apium graveolens ), caused by the fungus Septoria apiicola , is a major disease for which there is only limited host resistance. Some factors affecting the response of the fungus to essential oil components of host tissue were investigated using a susceptible blanching celery (var. Celebrity) and a more resistant green celery (var. Cutting) as hosts. After infection the time taken for pycnidia to appear on the surface was 16 days in Cutting and 13 days in Celebrity, but only limited leaf necrosis occurred in Cutting whereas the whole leaf rapidly became necrotic in Celebrity. Although the composition of the essential oil (limonene, β-pinene, terpinene, caryophyllene, sesquiterpenes and phthallides) was comparable in both varieties, Cutting contained over four times the total concentration of oil found in Celebrity. Extracts of the essential oil from Cutting were found to be more inhibitory to the growth of Septoria in culture. Of the terpenes tested (limonene, β-pinene, and caryophyllene), β-pinene was most inhibitory to the growth of hyphae on solid and liquid media. The role of the essential oil as part of a constitutive defense mechanism in host resistance to Septoria is discussed.     

The effect of powdery mildew (Erysiphe graminis f. sp. hordei) and leaf rust (Puccinia hordei) on spring barley in New Zealand. II. Apical development and yield potential

Reduced yields caused by powdery mildew and leaf rust in two seasons were associated with reduced plant growth. Combinations of early, late and full epidemics in one season, and 12 epidemic combinations in the second, were designed to identify crop sensitivity to disease by comparing growth and development with healthy plants. Early epidemics reduced ear number by increasing tiller death, and reduced grain number by effects on spikelet, floret or grain abortion, depending on the type of epidemic. Epidemics later in crop growth increased floret and grain abortion and also reduced grain weight.
There was no compensation by later-determined components for reduced growth and delayed development at earlier growth stages. Plants infected at early growth stages were more sensitive to late infections, seen as effects on the later-determined components, than plants which were healthy initially. Interactions occurred between epidemics at different times and are likely to occur between diseases and other constraints.