Intrapathotype diversity for aggressiveness and pathogen evolution in cultivar mixtures

ABSTRACT A model was developed and used to study the consequences of diversity for aggressiveness within pathotypes on pathogen evolution in two-component and four-component cultivar mixtures. It was assumed that, within a pathotype, a proportion of the isolates would have higher or lower spore efficacy than the average on a given host genetic background. Two situations were examined in which the pathogen can have either independent or negatively correlated values for spore efficacy on different cultivars. In the latter case, a pathogen genotype more aggressive than the average on a host genotype was always less aggressive on other host genotypes. In the simulations, isolates with greater aggressiveness relative to a host genotype were selected for and increased in frequency. However, because simple pathotypes always reproduced on the same host genotype whereas complex pathotypes were able to grow on several hosts, selection was faster for simple pathotypes. Pathotypes with two different levels of diversity for aggressiveness were compared with nondiversified pathotypes. In order to make comparisons, the effect of a 5 and 10% cost of virulence on the development of complex pathotypes was simulated. In general, increased diversity within pathotypes reduced the rate of increase of complex pathotypes in host mixtures, and this effect was stronger with greater frequencies of autodeposition of pathogen spores.     

Lesion expansion as an epidemic component

ABSTRACT A simulator for the enlargement of cohorts of circular lesions on cohorts of host tissue was used to examine five epidemiological parameters: radial rate (mm day(-1)) of lesion expansion, k (exp); maximum basic infection rate, R (m); proportion of lesion area as infectious, f; initial lesion size (mm(2)), z; and proportion of susceptible host sites, s. Based on the proportion of disease severity at day 50 and the proportion of the total disease that originated solely from lesion expansion, k(exp) was the most sensitive of the five parameters. A radial rate of only 0.1 mm day(-1) resulted in a proportion of >0.7 of the diseased area that came from lesion expansion. In an extensive survey of phytopathological literature, many plant pathogens had radial rates greater than 0.1 mm day(-1), which would result in a proportion of >0.95 of the total disease that comes from lesion expansion. Susceptible host sites, s, was a sensitive parameter, as this determined the host area into which lesions could expand. Naturally, R(m) was a sensitive parameter for the proportion of disease on day 50, as it controlled the overall speed of the epidemic. Initial lesion size was a relatively insensitive parameter, although z interacted significantly with s. The greatest proportion of disease that originated from lesion expansion occurred with fast k(exp), small z, and low values of s, R(m), and f. The model was validated with lesion numbers and severities obtained in natural epidemics of Cercospora medicaginis on alfalfa and Exserohilum turcicum on maize. We recommend that the 'epidemic quintuplet' used to describe polycyclic epidemics be expanded to the 'epidemic sextuplet' with the inclusion of k(exp), since lesion expansion is a major component of many polycyclic epidemics.     

Velocity of spread of wheat stripe rust epidemics

ABSTRACT Controversy has long existed over whether plant disease epidemics spread with constant or with increasing velocity. We conducted largescale field experiments with wheat stripe rust at Madras and Hermiston, Oregon, where natural stripe rust epidemics were rare, to test these competing models. Data from three location-years were available for analysis. A susceptible winter wheat cultivar was planted in pure stand and also in a 1:4 or 1:1 mixture with a cultivar immune to the stripe rust race utilized in the experiments. Plots were 6.1 m wide and varied from 73 to 171 m in length. A 1.5 by 1.5-m focus was inoculated in either the center (2001) or upwind of the center (2002 and 2003) of each plot. Disease severity was evaluated weekly throughout the epidemics in each plot at the same points along a transect running upwind and downwind from the focus. Velocity of spread was calculated from the severity data and regressed separately on time and on distance from the focus. In all location-years and treatments, and at all levels of disease severity, velocity consistently increased linearly with distance, at an average rate of 0.59 m/week per m, and exponentially with time. Further, across epidemics there was a significant positive relationship between the apparent infection rate, r, and the rate of velocity increase in both space and time. These findings have important implications for plant diseases with a focal or partially focal character, and in particular for the effectiveness of ratereducing disease management strategies at different spatial scales.     

Induced resistance in host mixtures and its effect on disease control in computer-simulated epidemics

The epidemic simulator EPIMUL was modified and used to study how induced resistance affected the development of epidemics in host mixtures. In the model, induced resistance resulted from the interaction of host tissue with avirulent spores and caused a reduction in the efficacy of virulent spores deposited afterwards. We denned three parameters to describe induced resistance: the level of protection, defined as the magnitude of reduction in the virulent spore efficacy for infecting host tissue; the host surface area protected by an interaction with one avirulent spore; and the duration of protection of the host tissue, in days. In our simulations, induced resistance slowed the epidemics and gave better disease control in the mixtures, even if protection lasted for only 2 days. The disease reduction in the mixture attributable to induced resistance was approximately proportional to the level of protection. The effect of induced resistance increased as the protected area increased. Epidemics were virtually unaffected by induced resistance restricted to the infection site, but the effect of induced resistance initially increased rapidly as larger areas were protected. There was little further gain as the protected area increased from 2·6% to 26%. The influence of induced resistance was reduced when the interactions between virulent and avirulent pathogens were reduced.     

Host diversity can reduce potato late blight severity for focal and general patterns of primary inoculum

ABSTRACT The use of host diversity as a tool for management of potato late blight has not been viewed as promising in the past. But the increasing importance of late blight internationally has brought new consideration to all potential management tools. We studied the effect of host diversity on epidemics of potato late blight in Oregon, where there was little outside inoculum. The experimental system consisted of susceptible potato cv. Red LaSoda and a highly resistant breeding selection, inoculated with local isolates of US-8 Phytophthora infestans. Potatoes were grown in single-genotype plots and also in a mixture of 10 susceptible and 26 resistant potato plants. Half of the plots received inoculation evenly throughout the plot (general inoculation) and half received an equal quantity of inoculum in only one corner of the plot (focal inoculation). The area under the disease progress curve (AUDPC) was greater in single genotype stands of susceptible cv. Red LaSoda inoculated throughout the plot than with stands inoculated in one focus. The host-diversity effect on foliar late blight was significant in both years of the investigation; the AUDPC was reduced by an average of 37% in 1997 and 36% in 1998, compared with the mean disease level for the potato genotypes grown separately. Though the evidence for influence of inoculum pattern on host-diversity effects was weak (P = 0.15), in both years there was a trend toward greater host-diversity effects for general inoculation. Statistical significance of host-diversity effects on tuber yield and blight were found only in one of the two years. In that year, tuber yield from both the resistant and susceptible cultivar was increased in mixtures compared with single genotype stands and tuber blight was decreased in mixtures for susceptible cv. Red LaSoda.     

Effect of genotype unit number and spatial arrangement on severity of yellow rust in wheat cultivar mixtures

Pure stands of a yellow rust-susceptible wheat cultivar, pure stands of a resistant cultivar, and a 1 : 1 random mixture of resistant and susceptible cultivars were compared to populations in which strips or hills of the cultivars were alternated to attain genotype units (units of the same host genotype) that were larger in area than that of a single wheat plant. These four host populations were grown in plots of different sizes in order also to alter the number of units per host population. The goal was to determine if increasing the number of genotype units in mixed populations of large genotype units improved disease control relative to pure-line populations by increasing the amount of inoculum exchange among genotype units. Random mixtures of the two cultivars always provided better disease control than did alternating strips or hills. Evidence for an effect of genotype unit number on the efficacy of mixtures for rust control was found in only one of three experiments. Random mixtures of the two cultivars increased grain yield relative to the pure stand mean, but alternating strips did not.     

Effect of crop growth and canopy filtration on the dynamics of plant disease epidemics spread by aerially dispersed spores

Most mathematical models of plant disease epidemics ignore the growth and phenology of the host crop. Unfortunately, reports of disease development are often not accompanied by a simultaneous and commensurate evaluation of crop development. However, the time scale for increases in the leaf area of field crops is comparable to the time scale of epidemics. This simultaneous development of host and pathogen has many ramifications on the resulting plant disease epidemic. First, there is a simple dilution effect resulting from the introduction of new healthy leaf area with time. Often, measurements of disease levels are made pro rata (per unit of host leaf area or total root length or mass). Thus, host growth will reduce the apparent infection rate. A second, related effect, has to do with the so-called "correction factor," which accounts for inoculum falling on already infected tissue. This factor accounts for multiple infection and is given by the fraction of the host tissue that is susceptible to disease. As an epidemic develops, less and less tissue is open to infection and the initial exponential growth slows. Crop growth delays the impact of this limiting effect and, therefore, tends to increase the rate of disease progress. A third and often neglected effect arises when an increase in the density of susceptible host tissue results in a corresponding increase in the basic reproduction ratio, R(0), defined as the ratio of the total number of daughter lesions produced to the number of original mother lesions. This occurs when the transport efficiency of inoculum from infected to susceptible host is strongly dependent on the spatial density of plant tissue. Thus, crop growth may have a major impact on the development of plant disease epidemics occurring during the vegetative phase of crop growth. The effects that these crop growth-related factors have on plant disease epidemics spread by airborne spores are evaluated using mathematical models and their importance is discussed. In particular, plant disease epidemics initiated by the introduction of inoculum during this stage of development are shown to be relatively insensitive to the time at which inoculum is introduced.     

Epidemics of Diaporthe adunca in experimental and in natural populations of Plantago lanceolata and the effect of partial resistance on disease development

Epidemics of the splash-dispersed pathogenic fungus Diaporthe adunca on its host, the perennial herb Plantago lanceolata , were followed during two consecutive years in transects at roadsides in the Netherlands. Epidemics of D. adunca were also studied on clones of a susceptible and a partially resistant genotype of P. lanceolata grown either in a pure stand or in a 1:1 mixture in small plots in the garden. The epidemics in the natural and experimental populations could be adequately described by logistic and Gompertz models, but large differences were found in final disease levels and relative growth rates. The effect of partial resistance on the epidemic in the mixture was less than in a pure stand, probably due to the provision of inoculum from the highly diseased susceptible genotype to the partially resistant genotype. In the garden focal and wind-direction effects were seen. In the natural populations the epidemics developed from numerous primary infected scapes making foci and wind-directions effects less conspicuous.     

Development of natural late blight epidemics in pure and mixed plots of potato cultivars with different levels of partial resistance

Late blight, caused by Phytophthora infestans , is the most severe disease of potato worldwide. Controlling late blight epidemics is difficult, and resistance of host cultivars is either not effective enough, or too easily overcome by the pathogen to be used alone. In field trials conducted for 3 years under natural epidemics, late blight severity was significantly lower in a susceptible cultivar growing in rows alternating with partially resistant cultivars (mixtures) than in unmixed plots of the susceptible cultivar alone. Partially resistant cultivars behaved similarly in unmixed and mixed plots. Mixtures of cultivars reduced disease progress rates and sometimes delayed disease onset over unmixed plots, but did so significantly only for the slowest epidemic. This suggests that reduction of area under the disease progress curve (AUDPC) in mixtures resulted from the cumulative action of minor effects. Disease distribution was focal in all plots at all dates, as shown by Morisita's index values significantly exceeding 1. Significant yield increases for the susceptible cultivar, and occasionally for the partially resistant ones, were observed in mixed-cultivar plots compared with single-cultivar plots. These results show that cultivar mixtures can significantly reduce natural, polycyclic epidemics in broadleaved plants attacked by pathogens causing rapidly expanding lesions.     

模拟植物病害流行时间动态的通用模型——Richards函数

 本文介绍了Richards生长函数及其在植物病害流行时间动态模拟中的应用。该函数的微分形式为(dx)/(dt)=rx((1/x)1-m-1)/(1-m),式中r为病害发展的速率,x表t日期的病情值率(0 < x < 1),m为流行曲线的形状参数。当m=0,m→1,m=2和m→∞时,从理论上证明Richards函数成为单分子、Compertz、Logistic和指数函数模型。以水稻纹枯病和马铃薯晚疫病的田间进展曲线进行摸拟分析发现,当m取值适当时还可获得较Gompertz或Logistic更逼真的Richards病害曲线拟合模型,而适当m取值的Richards模型比小分子模型对玉米粗缩病的拟合性也更好。因此认为,Richards函数是植病流行时间动态的通用模型。     

Allocation of resources: sources of variation in fusarium head blight screening nurseries

ABSTRACT Severe epidemics of Fusarium head blight (FHB) caused by Fusarium graminearum, group II (teleomorph: Gibberella zeae) have been occurring on wheat crops in the northcentral United States and southern Canada. Evaluation of resistance to FHB is difficul, because resistance is partial and infection depends upon host plant maturity. Variance component analysis was conducted to determine how best to allocate resources among environments, replications, and subsamples (heads per plot) in FHB screening nurseries. Advanced breeding lines from the Ohio State University wheat-breeding program were evaluated in screening nurseries from 1995 to 1997. Nurseries were artificially inoculated and sprinkler-irrigated to induce FHB epidemics. Over 80% of the variation within an environment resulted from variation associated with subsampling individual heads within plots. The second greatest source of variation was due to genotype by replication interactions. Host plant maturity influenced disease ratings in 1997. The repeatability of genotype means was approximately 50% within environments. The greatest reduction in genotype standard errors was obtained through additional environments, and then replications. Because the cost of an additional environment was estimated at five times the cost of an additional replication, the most cost-effective improvement in precision was obtained through the addition of replications. Advanced breeding lines should be evaluated in at least four replications per environment. Segregating populations will require more replications.     

Spread of faba bean rust over a discontinuous field

Spatio-temporal progress of an epidemic of faba bean rust was monitored over a discontinuous field. Trap plots were sown at increasing distances from a source plot, in the centre of which plants were inoculated. Disease spread in the source plot followed a focal pattern, with a radial velocity of expansion slightly lower than 0.1 m per day. At the end of the experiment, all trap plots had been infected, and two of the most distant ones showed unexpected high disease severity. Using a three-dimension model of disease progress, we showed that the epidemics on the scales of the source plot and of the trap plots could not be combined into a single epidemic on the whole-field scale. The epidemics had equivalent infection rates on both scales, but changing the scale dramatically affected their distance parameter. The epidemic in the source plot could have been caused by a short-distance, high-frequency, deterministic mechanism of spore dispersal, whereas infection of the trap plots could have been caused by a long-distance, low-frequency stochastic mechanism of spore dispersal. Although our results agreed with the predictions of a simulation model postulating these two mechanisms, alternative hypotheses which could also explain the observed disease pattern remained to be tested.     

Effects of planting density and the composition of wheat cultivar mixtures on stripe rust: an analysis taking into account limits to the replication of controls

ABSTRACT The effect of plant density on disease is not well understood in populations of a single host plant genotype and has been studied even less in mixtures of host genotypes. We performed an experiment to evaluate the effect of wheat planting density on infection by Puccinia striiformis in experimental plots with a single wheat genotype and in plots with two genotypes making up a range of frequencies. Stripe rust severity in single-genotype plots increased with planting density in 1997 but decreased with planting density in 1998. Disease in host mixtures was compared to the weighted mean of disease levels in the corresponding single-genotype plots. The design of the field experiment included limited replication of these reference treatments (that is, there was not a unique pair of single-genotype plots for each mixture plot); therefore, we devised an analysis based on collapsing the data into independent mean observations. Disease reduction due to host diversity was less when one genotype predominated than when both host genotypes were present at nearly equal frequencies. The greatest mean host-diversity effect for reduced disease was at the intermediate planting density of 250 seeds per m(2).     

Host mixture efficacy in disease control: effects of lesion growth analyzed through computer-simulated epidemics

Lesion growth varies among foliar parasites and in order to study the effect of lesion growth on the efficacy of host mixtures to control epidemics, we altered the epidemic simulator Epimul by integrating a lesion growth function into the model. A theoretical study was performed by simulating epidemics caused by parasites with different lesion growth rates, spore dispersal gradients and multiplication rates. We found that increases in lesion growth rates resulted in large decreases in the effectiveness of mixtures for disease control and interacted strongly with parasite multiplication rate and spore dispersal gradient. The decline in mixture efficacy for epidemics with high lesion growth rates was reduced when parasite multiplication rate was higher and spore dispersal gradient steeper. Our results suggested that the lower number of infections on susceptible plants in the mixture as a result of inoculum loss on resistant hosts was partially compensated by lesion growth.     

Qualification of a Plant Disease Simulation Model: Performance of the LATEBLIGHT Model Across a Broad Range of Environments

ABSTRACT The concept of model qualification, i.e., discovering the domain over which a validated model may be properly used, was illustrated with LATEBLIGHT, a mathematical model that simulates the effect of weather, host growth and resistance, and fungicide use on asexual development and growth of Phytophthora infestans on potato foliage. Late blight epidemics from Ecuador, Mexico, Israel, and the United States involving 13 potato cultivars (32 epidemics in total) were compared with model predictions using graphical and statistical tests. Fungicides were not applied in any of the epidemics. For the simulations, a host resistance level was assigned to each cultivar based on general categories reported by local investigators. For eight cultivars, the model predictions fit the observed data. For four cultivars, the model predictions overestimated disease, likely due to inaccurate estimates of host resistance. Model predictions were inconsistent for one cultivar and for one location. It was concluded that the domain of applicability of LATEBLIGHT can be extended from the range of conditions in Peru for which it has been previously validated to those observed in this study. A sensitivity analysis showed that, within the range of values observed empirically, LATEBLIGHT is more sensitive to changes in variables related to initial inoculum and to weather than to changes in variables relating to host resistance.     

Resistance to early blight of tomato with respect to various parameters of disease epidemics

Resistance to early blight in the tomato was assessed by examining various parameters of the progress of the disease. Artificial inoculation and the scoring technique were standardized. Test plants were inoculated with 125cfu/ml of a 12-day-old culture of a pathogenic isolate of Alternaria solani. Screening under artificial conditions was more informative than that under natural epidemic conditions. Tomato cultivars CLN-2071-C, CLN-2070-A, BSS-174, and DTH-7 with resistance expressed as slow blighting against four pathogenic isolates of A. solani, were selected for cultivation in disease-prone areas. Disease intensity increased with the age of plants under the same inoculum load. The area under the disease progress curve (AUDPC) was positively correlated with the percentage disease index and negatively with resistance. Calculation of the apparent infection rate (r) was more informative for natural epidemics than for artificial conditions. The sequential apparent infection rate between observation periods was better correlated with disease progress than was the total apparent infection rate between the first and last observations. A double sigmoidal disease progress curve during the same cropping season was characteristic of some varieties when fungal infection took place during the vegetative phase of crop growth.     

Effects of moisture and septoria tritici blotch stresses on wheat yields under semi-arid conditions: A Simulation study

The effects of Septoria tritici blotch on spring wheat were incorporated into a crop growth simulator and the model was then used to evaluate the interrelationships between moisture and disease stresses on wheat yields under semi-arid conditions. Simulation experiments revealed that moisture availability plays a role in determining the relationship between disease and yield. In a season when precipitation was distributed evenly, a non-diseased crop yielded more than diseased crops. Yield reductions in diseased crops were related to the severity of epidemics; the relationship between disease and yield was inverse and linear. In a season when water was scarce during the stage of kernel-filling, crops with mild or moderate disease epidemics (apparent infection rate of a logistic model,r, of 0.025 or 0.05 per unit) yielded more than the non-diseased crop. The yield of the crop with a severe epidemic (r of 0.1 per unit) was reduced substantially and the relationship between disease and yield was parabolic. The parabolic relationship was linearized when additional precipitation was simulated in the model.     

The effects of dispersal gradient and pathogen life cycle components on epidemic velocity in computer simulations

ABSTRACT The velocity of expansion of focal epidemics was studied using an updated version of the simulation model EPIMUL, with model parameters relevant to wheat stripe rust. The modified power law, the exponential model, and Lambert's general model were fit to primary disease gradient data from an artificially initiated field epidemic of stripe rust and employed to describe dispersal in simulations. The exponential model, which fit the field data poorly (R (2) = 0.728 to 0.776), yielded an epidemic that expanded as a traveling wave (i.e., at a constant velocity), after an initial buildup period. Both the modified power law and the Lambert model fit the field data well (R(2) = 0.962 to 0.988) and resulted in dispersive epidemic waves (velocities increased over time for the entire course of the epidemic). The field epidemic also expanded as a dispersive wave. Using parameters based on the field epidemic and modified power law dispersal as a baseline, life cycle components of the pathogen (lesion growth rate, latent period, infectious period, and multiplication rate) and dispersal gradient steepness were varied within biologically reasonable ranges for this disease to test their effect on dispersive wave epidemics. All components but the infectious period had a strong influence on epidemic velocity, but none changed the general pattern of velocity increasing over time.     

Effect of wheat cultivar mixtures on populations of Puccinia striiformis races*

This study quantifies the frequency of simple and complex races (races that can infect two or more components) of Puccinia striiformis in mixtures of wheat cultivars possessing different race-specific resistance genes. Treatments were designed so that the complex race changed depending on the host mixture, thus enabling us to observe the influence of pathogen complexity in different genetic backgrounds. Six cultivar mixtures and one pure stand of winter wheat were inoculated with three races of P. striiformis at two locations for two seasons. Potted plants of three winter wheat cultivars, each susceptible to one of the three races of the pathogen, were used to sample the pathogen during the field epidemics. Disease incidence on the differential cultivars was used to calculate the proportion of the three races in each treatment. The specific cultivars included in the mixtures influenced the frequencies of the three races. Increasing the number of virulent races in a mixture reduced the frequency of the complex race relative to the other two races. The results suggest that genetic background of the pathogen race, host composition, and interaction among pathogen races may be as important as cost of virulence in determining race frequencies in mixtures.     

Characterization of early leaf spot suppression by strip tillage in peanut

ABSTRACT Epidemics of early leaf spot of peanut (Arachis hypogaea), caused by Cercospora arachidicola, are less severe in strip-tilled than conventionally tilled fields. Experiments were carried out to characterize the effect of strip tillage on early leaf spot epidemics and identify the primary target of suppression using a comparative epidemiology approach. Leaf spot intensity was assessed weekly as percent incidence or with the Florida 1-to-10 severity scale in peanut plots that were conventionally or strip tilled. The logistic model, fit to disease progress data, was used to estimate initial disease (y(0)) and epidemic rate (r) parameters. Environmental variables, inoculum abundance, and field host resistance were assessed independently. For experiments combined, estimated y(0) was less in strip-tilled than conventionally tilled plots, and r was comparable. The epidemic was delayed in strip-tilled plots by an average of 5.7 and 11.7 days based on incidence and severity, respectively. Tillage did not consistently affect mean canopy temperature, relative humidity, or frequency of environmental records favorable for infection or spore dispersal. Host response to infection was not affected by tillage, but infections were detected earlier and at higher frequencies with noninoculated detached leaves from conventionally tilled plots. These data suggest that strip tillage delays early leaf spot epidemics due to fewer initial infections; most likely a consequence of less inoculum being dispersed to peanut leaves from overwintering stroma in the soil.