Patterns of development of Septoria nodorum and S. tritici in some winter wheat crops in Western Europe, 1981-83

In data collected at 19 sites in Western Europe during 1981-83. two patterns of development of Septoria nodorum and S. tritici on foliage of winter wheal were distinguished. In sudden outbreaks, lesions appeared simultaneously on the upper leaf layers of crops, usually after the end of stem extension; these outbreaks were ascribed to short, heavy rain storms in which pycnidiospore inoculum in basal leaves was elevated up to 60 cm through the crop canopy. Gradual epidemics were characterized by disease arising on successive leaf layers as they appeared during sustained periods of weather suitable for inoculum transport and infection.
The data indicate incubation periods of 2-4 weeks for S. nodorum and 3-5 weeks for S. tritici. it is suggested that a leaf layer cannot normally sustain more than one pathogen generation and that its infection arises from inoculum borne on leaves older than in the layer situated immediately below it. The potential level of disease in a crop may relate to the amount of inoculum present in spring. The proportions of disease caused by the two Septoria species varied greatly between sites and years, but the data provided no explanation.
It is concluded that a septoria forecast scheme needs to recognise the importance of sudden disease outbreaks and to include not only weather but also host growth and inoculum factors.     

A dynamic simulation model for powdery mildew epidemics on winter wheat*

A system dynamic model for epidemics of Blumeria graminis (powdery mildew) on wheat was elaborated, based on the interaction between stages of the disease cycle, weather conditions and host characteristics. The model simulates the progress of disease severity, expressed as a percentage of powdered leaf area, on individual leaves, with a time step of one day, as a result of two processes: the growth of fungal colonies already present on the leaves and the appearance of new colonies. By means of mathematical equations, air temperature, vapour pressure deficit, rainfall and wind are used to calculate incubation, latency and sporulation periods, the growth of pathogen colonies, infection and spore survival. Effects of host susceptibility to infection, and of leaf position within the plant canopy, are also included. Model validation was carried out by comparing model outputs with the dynamics of epidemics observed on winter wheat grown at several locations in northern Italy (1991–98). Simulations were performed using meteorological data measured in standard meteorological stations. As there was good agreement between model outputs and actual disease severity, the model can be considered a satisfactory simulator of the effect of environmental conditions on the progress of powdery mildew epidemics.     

Influence of crop growth and structure on the risk of epidemics by Mycosphaerella graminicola (Septoria tritici) in winter wheat

Generally, it is recognized that inocula of Septoria tritici present on the basal leaves of winter wheat crops are spread towards the top of the canopy by splashy rainfall. This mechanism of inoculum dispersal is commonly accepted to be a key limit on disease progression. Therefore, attempts to forecast epidemics of S. tritici often quantify rainfall by some means, but largely ignore measurement of pathogen and host variables. In the present study, we show that new wheat leaves emerge initially at a height below established leaves that can contain sporulating lesions of S. tritici . This presents the possibility of horizontal inoculum transfer, even without splashy rainfall. The extent and duration of overlap between emergent and established leaves was found to differ considerably with cultivar and sowing date. Nitrogen application had little effect on overlap, because differences in crop phenology, e.g. leaf area and nodal length, were relative. However, estimates of raindrop penetration to the base of crop canopies suggested that vertical movement of inoculum is affected by nitrogen application. Crops receiving more nitrogen are denser, and therefore less rainfall reaches the base of the canopy. The interactions between crop and pathogen development are discussed with reference to the implications for predicting disease risk. In particular, cultivar traits that promote disease escape are quantified.     

Factors determining the severity of epidemics of Mycosphaerella graminicola (Septoria tritici) on winter wheat in the UK

Epidemics of disease caused by Septoria tritici were studied in detail in 11 crops of winter wheat cv. Longbow over 4 years. Serious damage to the uppermost two leaf layers was caused by splash-borne infection from lower in the crop early in the life of the leaves, followed by one or rarely two cycles of multiplication within a leaf layer. Infection conditions rarely limited damage, even in a dry year; the timing and, to a lesser extent, amount of initial inoculum movement to an upper leaf layer was of greater importance. Timing of initial infection was determined by when rain splash occurred in relation to emergence of a leaf layer. Occurrence of infections soon after a leaf layer started to emerge allowed more time for multiplication of disease within that layer. These infections tended to be more severe because the leaves were closer to inoculum sources within the crop. Slight differences in phenology between locations explain why initially random disease distributions sometimes become aggregated. Early-sown crops are at greater risk because they mature more slowly, allowing more disease multiplication and better transfer between leaf layers.     

An epidemiological simulation model with three scales of spatial hierarchy

ABSTRACT An epidemiological model integrating three organizational scales of host plant populations (e.g., sites, leaves, and plants) is presented. At the lowest (site) scale, the model simulates the dynamics of vacant, latent, infectious, and removed sites. Three types of vacant sites are distinguished, depending on presence of infections at higher scales (leaf or plant). The rate of infection of each type of vacant site is computed according to ratios of autodeposition, allo-leaf-deposition, and allo-plantdeposition. At the leaf and plant scales, the rate of victimization is a function of the rate of infection of vacant sites. Sensitivity analyses showed that deposition patterns (the relative proportions of auto-, allo-leaf-, and allo-plant-depositions) and host structure (leaf size and number of leaves per plant) affected the speed of epidemics at the different scales. Model outputs conformed with results from other approaches in the case of random distribution of the disease. The model hypotheses concerning infection from autodeposited propagules, and their implications for disease epidemics, are discussed. The model can be used to derive relationships between allo-deposition ratios and disease incidences at the three scales. These relationships become simple when disease intensity is low. These relationships may be useful, e.g., to assess the potential efficiency of cultivar mixture to control epidemics. Integration of different organization scales and allo-deposition parameters enables the model to capture important features of epidemics developing in space without using explicitly spatialized variables. Such an approach could be useful to analyze other ecological processes that involve a variety of scales.     

Effect of Fungicides on Septoria nodorum and Wheat Green Leaf Area1

Fungicides applied at different growth stages were compared for the control of wheat glume blotch, caused by Septoria nodorum (Berk.) Berk. In glasshouse and small-scale field tests, 11 commercially used fungicides, when applied to fully emerged wheat ears, cv. Hobbit, gave only partial control of ear infection by spores applied immediately after spraying. They gave better control of infection of leaves by a similar inoculum. Larger field trials, moreover, indicated better foliar and ear disease control from fungicide applications made before ear emergence, compared with those applied later. In another field trial, nine fungicides were applied to replicated plots of winter wheat, cultivars Atou and Hobbit at GS 31, GS 39, GS 55 or GS 71. On five occasions from anthesis (25 June) until 30 July disease was assessed and green leaf area measured. The trial was harvested on 22 August 1980. Significantly greater yields and grain weights were obtained from plots sprayed with fungicide at GS 39 than from those sprayed at any other time or left unsprayed. Treatments made before symptoms of infection by S. nodorum developed (i.e. before GS 55) prolonged the duration of green leaf areas during the post-anthesis period. Grain weight and yield increases were related closely to the duration of green leaf area above 75 96, but not to the amount of green leaf area at any single assessment nor to the subsequent degree of disease development. The data on which this abstract is based form part of a continuing research programme, the results of which will be published in full at a later date. Full details on the 1980 results referred to above appear in the Annual Report of Long Ashton Research Station (1980).     

Modeling of Relationships Between Weather and Septoria tritici Epidemics on Winter Wheat: A Critical Approach

ABSTRACT Two models for predicting Septoria tritici on winter wheat (cv. Riband) were developed using a program based on an iterative search of correlations between disease severity and weather. Data from four consecutive cropping seasons (1993/94 until 1996/97) at nine sites throughout England were used. A qualitative model predicted the presence or absence of Septoria tritici (at a 5% severity threshold within the top three leaf layers) using winter temperature (January/February) and wind speed to about the first node detectable growth stage. For sites above the disease threshold, a quantitative model predicted severity of Septoria tritici using rainfall during stem elongation. A test statistic was derived to test the validity of the iterative search used to obtain both models. This statistic was used in combination with bootstrap analyses in which the search program was rerun using weather data from previous years, therefore uncorrelated with the disease data, to investigate how likely correlations such as the ones found in our models would have been in the absence of genuine relationships.     

The use of rainfall and accumulated mean temperature to indicate fungicide activity in the control of leaf diseases of winter wheat in the UK1

During 1988/1990, a series of 21 experiments was established in commercial crops of winter wheat. Chlorothalonil, fenpropimorph and propiconazole were chosen as protectant, eradicant or curative fungicides active against leaf diseases of winter wheat in the UK. To test their properties each one was applied once only to separate plots during a period of 7–8 consecutive weeks in May and June (GS 32–39). Disease progress was assessed weekly on adjacent unsprayed control and sprayed plots up to GS 85. Septoria tritici leaf blotch (Mycosphaerella graminicola) was the disease that occurred most frequently and severely across the 21 sites. Powdery mildew (Erysiphe graminis), brown rust (Puccinia recondita) and yellow rust (P. striiformis) occurred at fewer sites and were sufficiently severe to distinguish differences between the active ingredients at only two or three sites. Analysis of the disease progress curves for 75% control of each disease at one site only indicated that chlorothalonil possessed very good protectant but shorter-term eradicant activity against M. graminicola and P. striiformis. Fenpropimorph exhibited only short-term eradicant control of M. graminicola, but gave excellent protectant and eradicant control of E. graminis and P. striiformis; against P. recondita only eradicant activity was apparent. Propiconazole showed activity similar to that of fenpropimorph against P. recondita and excellent protectant and eradicant activity against M. graminicola and P. striiformis; against E. graminis, it gave good protectant and eradicant control. From disease progress curves, it was possible to calculate the period of protectant and eradicant activity in thermal time (accumulated degree days above zero) for each of the three active ingredients and to identify the most effective timing(s) for fungicide application in relation to rainfall or imputed infection.     

A dynamic model forecasting infection of pear leaves by conidia of <Emphasis Type="Italic">Venturia nashicola</Emphasis> and its evaluation in unsprayed orchards

A dynamic model, called VenInf, was developed to forecast infection of pear leaves by conidia of Venturia nashicola. By simulating conidial infection processes following a rain event, the model estimates % conidia that successfully infected leaves at the end of an infection period. The model is mainly derived from logistic models developed from recent laboratory and glasshouse experimental results on infection of pear seedlings to estimate the rates of infection and mortality. It simulates the conidial infection process at 5 min intervals using temperature, relative humidity (RH), surface wetness and rainfall as input. The model was evaluated against pear scab in four unsprayed orchards in China over a 4-year period. In all orchards, all significant disease increases were associated with infection periods predicted by the model. In one orchard, in 2004 the incidence of leaf infection remained very low (<3%) during the entire season despite the model forecasting several severe infection periods. Results of orchard evaluation suggest that the model is able to identify all important potential infection periods. Thus, further field studies should be carried out to determine whether and how the model can be used in practice to assist farmers in making decisions on fungicide applications.     

The role of temperature and rainfall in the epidemiology of Puccinia striiformis f.sp. tritici in the summer rainfall area of eastern Australia

Infection of wheat seedlings by Puccinia striiformis f.sp. tritici was investigated under both laboratory (constant temperature) and field conditions using a 15-h period of 100% r.h. In laboratory studies, infection decreased from 100% at 15 4 C to 0.8% at 20.5 C, and it was estimated that no infection would occur at or above 20.8 + 0.2 C. In contrast, high levels of infection occurred under field conditions even when temperatures fluctuated within the range 19–30°C. Overnight infection experiments conducted at Toowoomba over an 18-month period demonstrated that periods of moisture and temperature favouring infection by P. striiformis f.sp. tritici occurred regularly, even during summer, and that high temperature was a limiting factor on only 3% of the nights tested. This indicates that stripe rust could oversummer readily in this region of the eastern Australian wheat belt, at least in sheltered areas, given a susceptible host. Regression equations relating mean temperature to infection and minimum temperature to infection identified mid to late autumn as an important period in the epidemic development of the disease in this region. The amount of rain recorded during this period was closely associated with subsequent levels of stripe rust observed in commercial crops over the years 1983–87.     

Rain-driven epidemics ofPhytophthora porri on leek

White tip disease of leek (Allium porrum), caused byPhytophthora porri, was studied in field experiments. On fields infested by soil-borne inoculum (oospores), relatively short periods of explosive disease increase alternated with periods in which apparently no new infections occurred. The analysis of rain data and disease data, using a degree-day model for incubation periods at constant temperatures, confirmed the hypothesis that disease increase ofP. porri is significantly correlated with rain; R _adj ² was 0.91, 0.41 and 0.51 in 1992, 1993 and 1994, respectively. Correlations were highest early in the season. Lack of correlation later in the season may be ascribed to the effect of lesion death, which may be caused by total or partial leaf death, by desiccation or by other fungi overgrowingP. porri, and to the effect of secondary infection by zoosporangia, which appears to be not so strongly rain-driven as primary infection. Zoosporangia were observed in fields on water-logged light-green lesions. High lesion densities of leaf tips and leaf units at 10–20 cm above the leaf axils indicated that most infections depend on free water, either in puddles or in a water basin near the leaf axils. Although disease correlates well with rain data, disease forecasts will be unreliable as long as rain forecasts are unreliable.     

The role of rain in dispersal of the primary inoculum of Plasmopara viticola

Although primary infection of grapevines by Plasmopara viticola requires splash dispersal of inoculum from soil to leaves, little is known about the role of rain in primary inoculum dispersal. Distribution of rain splashes from soil to grapevine canopy was evaluated over 20 rain periods (0.2 to 64.2 mm of rain) with splash samplers placed within the canopy. Samplers at 40, 80, and 140 cm above the soil caught 4.4, 0.03, and 0.003 drops/cm(2) of sampler area, respectively. Drops caught at 40 and 80 cm (1.5 cm in diameter) were larger than drops at 140 cm (1.3 cm). Leaf coverage by splashed drops, total drop number, and drop size increased with an increase in the maximum intensity of rain (mm/h) during any rain period. Any rainfall led to infection in potted grapevines placed outside on leaf litter containing oospores, if the litter contained germinated oospores at the time of rain; infection severity was unrelated to rain amount or intensity. Results from vineyards also indicate that any rain can carry P. viticola inoculum from soil to leaves and should be considered a splash event in disease prediction systems. Sampling for early disease detection should focus on the lower canopy, where the probability of splash impact is greatest.     

Effects of simulated acid rain and root-knot nematode on tomato

Effects were examined of simulated acid rain and of root-knot nematode, Meloidogyne incognita race 1, on plant growth, yield, photosynthetic pigments and leaf epidermal characters of tomato ( Lycopersicon esculentum cv. Pusa Ruby). Sequential inoculation exposures (pre-, post-, and concomitant with nematode inoculation) were carried out in a greenhouse. Intermittent treatments of simulated acid rain at pH 3.2 caused white-to-tan irregular lesions on both the upper and lower surfaces of tomato leaves. The foliar symptoms were more pronounced on nematode-infected plants. Simulated acid rain (pH 3.2) and/or nematode infection suppressed plant growth, yield and pigment synthesis, the effects being greatest in post-inoculation treatments compared with simulated acid rain at pH 6.8. The total weight of fruits per plant was greatly suppressed owing to simulated acid rain or nematode infection. Chlorophyll a was found to be more sensitive to simulated acid rain or nematode infection than other leaf pigments. Root penetration, galling, egg mass production, and fecundity (number of eggs per egg mass) of M. incognita were enhanced at pH 5.6 and suppressed at pH 3.2 compared with pH 6.8. Nematode infection or simulated acid rain at pH 3.2 suppressed stomata and trichome development (number and size). Simulated acid rain treatments at pH 5.6 had a positive effect on number and size of trichomes, but a negative effect on stomata. The apertures of stomata were wider on tomato leaves exposed to simulated acid rain, especially at pH 3.2, than at pH 6.8.     

Effects of Pyrenophora teres on dry matter production and yield components of winter barley

Inoculation of winter barley plants in a glasshouse with Pyrenophora teres at three different growth stages (GS 11, 13 or 30) greatly decreased root and shoot dry matter production and the size of healthy leaves produced subsequently, but there was no significant yield loss with a single inoculation at GS 11 or 30. Inoculation at GS 13, GS 31 or GS 39 decreased grain yield by 12,17 and 20% respectively, and also straw yield. Larger yield losses resulted from repeated inoculations on five successive occasions (GS 11, 13, 30, 31 and 39) which caused much disease on all leaves throughout the life of the plant. All components of yield measured were decreased by the cumulative effect of successive inoculations; ear number by 15%), grain sites per ear by 20%, grain yield by 48% and straw yield by 32%.     

Modelling the progress of light leaf spot (Pyrenopeziza brassicae) on winter oilseed rape (Brassica napus) in relation to leaf wetness and temperature

A compartmental model was developed to describe the progress with time of light leaf spot ( Pyrenopeziza brassicae ) on leaves of winter oilseed rape ( Brassica napus ) during the autumn in the UK. Differential equations described the transition between the four compartments: healthy susceptible leaves, infected symptomless leaves, sporulating symptomless leaves and leaves with necrotic light leaf spot lesions, respectively. The model was fitted to data on the progress of light leaf spot on winter oilseed rape at a single site during the autumn of the 1990–1991 season. Model parameters were used to describe rates of leaf appearance, leaf death, infection by airborne ascospores (primary inoculum) and infection by splash-dispersed conidiospores (secondary inoculum). Infection was dependent on sufficient leaf wetness duration. The model also included delay terms for the latent period between infection and sporulation and the incubation period between infection and the appearance of necrotic light leaf spot lesions. This modified SEIR model formulation gave a reasonable fit to the experimental data. Sensitivity analysis showed that varying the parameter accounting for the rate of infection by ascospores affected the magnitude of the curves after the start of the epidemic, whilst including a parameter for conidiospore infection improved the fit to the data. Use of ascospore counts from different sites and different years showed variation in spore release patterns sufficient to affect model predictions.     

Effects of leaf wetness duration, relative humidity, light and dark on infection and sporulation by Didymella rabiei on chickpea

No infection occurred at less than 95% relative humidity (r.h.) when chickpea plants were dried after inoculation with conidia of Didymella rabiei. Infection was significant when the dry leaves were exposed to 98% r.h. for 48 h. When inoculated plants were subjected to different leaf wetness periods, some infection occurred with 4 h wetness, and disease severity increased with wetness duration according to an exponential asymptote, with a maximum value after about 18 h. Germination of conidia and germ tube penetration increased linearly with increasing wetness periods when recorded 42 h after inoculation. With a 24-h wetness period, germination of conidia was first observed 12 h after inoculation and increased linearly with time up to 52 h (end of the experiment). Dry periods immediately after inoculation, followed by 24-h leaf wetness, reduced disease severity; as the dry period increased the severity decreased. Disease severity increased with increasing periods of darkness after inoculation. The number of pycnidia and the production of conidia on infected leaves increased only slightly with high r.h. (either in the light or in the dark), but large increases occurred over an 8-day period when the leaves were kept wet.     

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.     

Spatial distributions of Septoria nodorum and S. tritici within crops of winter wheat

In some crops of winter wheat selected from a range monitored in Western Europe during 1981-83, Septoria nodovum and S . tritici were spatially very uniformly distributed from the beginning of the growing season onwards. A cube root transformation produced a constant variance in lesion numbers per leaf, similar for both pathogens at about 0.5. This permits the sample size needed for a given accuracy to be estimated. Counts of conidia washed from leaves by a standard procedure had a constant coefficient of variation, independent of disease level. Large samples would be necessary for accurate counts, particularly if leaves from different layers were examined separately. The pattern of lesion numbers on leaves was best described by a negative binomial distribution: this predicts an incidence-severity curve to which the data conformed closely. Hence incidence estimates can be used to estimate severity, which may be more economical of sampling effort. Correlations between lesion counts on different leaves of the same tiller were negative and highly significant for S. nodorum in 1982, but positive and significant for S. tritici in 1983. The causes of this difference are unknown.     

Entwicklung einer Herbstprognose für den Erreger der Wurzelhals- und Stängelfaule Phoma lingam (Teleomorph: Leptosphaeria maculans)

Based on experimental data of oilseed rape monitoring in Schleswig-Holstein during the last seven years and actually established monitoring at four sites in Germany, a forecasting system for a determined autumn application against the important rape pathogen root collar (Phoma lingam; teleomorph: Leptosphaeria maculans) was developed. Early primary infections by ascospores in autumn are the elicitors of severe yield-reducing root collar epidemics, which depend on weather conditions, especially rainfall, leaf wetness and precipitation. Based on measured data and on trial weather records, a model was developed to predict the daily infection risk. Threshold values for a determined application were calculated based on disease incidence monitoring values. Applications predetermined from the calculated “daily infection value” reduced pathogen populations in autumn in comparison to the untreated control. Effects of the autumn treatments on root collar could be recorded; however, a significant yield effect occurred at one site only. Weather conditions over winter until vegetation start in the following year influence root collar severely. Due to this conclusion it is difficult to evaluate prosperous autumn treatments.     

The effect of powdery mildew (Erysiphe graminis f. sp. hordei) and leaf rust (Puccinia hordei) on spring barley in New Zealand. I. Epidemic development, green leaf area and yield

Powdery mildew and leaf rust caused large yield losses in spring barley grown near Christchurch, New-Zealand, in two seasons. Disease present during early growth stages was as damaging to yield as disease late in the season. Moderate leaf rust severities after anthesis were most damaging when combined with earlier mildew epidemics. Later growth did not compensate for reduced yield potential induced by early infection. This was attributed, at least in part, to an effect on leaf size, and therefore on green leaf area, at later growth stages. There was a closer relationship, by regression analysis, of yield to green leaf area than to disease severity in three cultivars.
The three cultivars. which differed in yield potential and disease resistance, were not equally sensitive to disease. It is proposed that high yielding cultivars may be the most sensitive to yield constraint by disease.