Developing smarter host mixtures to control plant disease

Adaptation of plant pathogens to disease control measures (both chemical and genetic) is facilitated by the genetic uniformity underlying modern agroecosystems. One path to sustainable disease control lies in increasing genetic diversity at the field scale by using genetically diverse host mixtures. In this study, a robust population dynamics approach was used to model how host mixtures could improve disease control. It was found that when pathogens exhibit host specialization, the overall disease severity decreases with the number of components in the mixture; this finding makes it possible to determine an optimal number of components to use. In a simple case, where two host varieties are exposed to two host‐specialized pathogen species or strains, quantitative criteria for optimal mixing ratios are determined. Using these model outcomes, ways to optimize the use of host mixtures to decrease disease in agroecosystems are proposed.     

Increasing complexity of resistance in host populations through intermating to manage rust of pearl millet

ABSTRACT Pearl millet inbreds Tift 23DB, Tift 85DB, PS748BC, and Tift 89D(2) were used to develop three categories of host mixtures (physical mixtures, random-mated populations, and mixtures of two-way and three-way crosses) representing different levels of complexity of resistance through increased heterogeneity within populations and through stacking of resistance genes within the heterogeneous populations. The potential of these mixtures to reduce rust epidemics was evaluated in the field. Area under the disease progress curves (AUDPCs) of all physical mixtures were less than the mean of the components in 1995 and were less than the mean of the components for five of the six mixtures in 1997. In 1996, AUDPCs of the physical mixtures were consistently greater than the mean of their components. AUDPCs of the random-mated mixtures and the mixtures of crosses were consistently less than the mean of the components in 1996 and 1997, with reductions ranging from 12 to 71%. Dry matter yield (DMY) of physical mixtures relative to the mean DMY of the components was inconsistent, ranging from 18% less to 50% more than the mean of the components. The random-mated populations and the mixtures of crosses yielded 18 to 40% more DMY than the mean yield of the pure stands of their components.     

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.     

The effect of variety mixtures on the development of swede powdery mildew

Infection by Erysiphe polygoni was followed on the components of five mixed populations containing different proportions of a partially resistant swede variety, Ruta Øtofte (RØ), and a highly susceptible variety, Ne Plus Ultra (NPU) (100:0; 75:25; 50:50; 25:75; 0:100). Disease levels remained much lower on RØ than on NPU in all populations. For several weeks, disease on NPU was greatest in pure plots and was progressively reduced in mixtures with increasing proportions of RØ. Later, disease levels on NPU were similar in all populations. Disease on RØ was not increased in mixtures with NPU. Root yield of NPU, but not of RØ, was greater in mixtures than in pure stands. The possible use of variety mixtures to control powdery mildew in swede crops is discussed.     

Competition and interactions among stripe rust pathotypes in wheat-cultivar mixtures

A series of experiments was conducted with wheat stripe rust to analyse competition between simple and complex pathotypes in host mixtures. Two different pathotype combinations were tested, with different host components. Each combination included a complex (able to infect two host components) and two simple pathotypes. For one of the combinations, induced resistance was tested in a separate experiment as a possible interaction among pathotypes. Disease severity and pathotype frequencies were measured three times during the epidemic, on each host component grown in pure stands and in mixtures. In one of the experiments, pathotype frequencies were also measured within secondary foci. One of the complex pathotypes appeared to have a low fitness on one of the host components and did not significantly increase in frequency in host mixtures relative to pure stands. The average frequency of the other complex pathotype increased during the first epidemic cycles, but remained stable afterwards, below expected values. The results suggest that the development of complex pathotypes in host mixtures may be influenced by differential aggressiveness on the host components, by induced resistance and by random effects resulting from the formation of disease foci, and depends on pathogen autoinfection rate and dispersal mechanisms.     

A simulation study on managing plant diseases by systematically altering spatial positions of cultivar mixture components between seasons

A simulation study was conducted to investigate epidemic development of a race‐specific pathogen in cultivar mixtures over six consecutive seasons where the spatial position of each mixture component was systematically altered between seasons. Results showed that, even for a relative large genotype unit area in a mixture, altering cultivar positions between seasons could, on average, increase disease suppression by a third over the corresponding mixture without position changes between seasons. Overall, the disease suppression achieved by mixtures with position change between seasons was close to that achieved by random mixtures. Greater redistribution distance of overwintered inoculum reduced the disease control efficacy achieved by change in the position of individual mixture components between seasons. It is therefore concluded that using mixtures with a relatively large genotype unit area together with systematic changes in the spatial positions of individual mixture components between seasons is a feasible option for integrated disease management.     

Assessing Four-Way Mixtures of Winter Wheat Cultivars from the Performances of their Two-Way and Individual Components

Seven four-way bread wheat mixtures were compared to their components (individual cultivars and two-way mixtures) for septoria tritici blotch severity, grain protein content, and grain yield. Four-way mixtures and two-way mixtures performed better than the average of individual pure line components. Disease severity and protein content were most influenced by mixtures, with mixture efficiencies being greater in the four-way than in the two-way mixtures. Performances of four-way mixtures in terms of diseased leaf area reduction, grain yield, 1000-grain weight, and grain protein content were better predicted by performances of two-way mixtures than by those of pure lines. Our results suggest that two-way mixtures should be screened to remove unfavourable cultivar pairs and those binary mixtures that show complementarity should be selected in order to construct four-way mixtures.     

Evolution of a pathogen population in host mixtures: simple race–complex race competition

The evolution of the genetic structure of a pathogen population was studied in a varietal mixture with an epidemic simulator based on the model EPIMUL. The pathogen population was composed of simple races able to develop on only one component of the mixture and a complex race which developed on all mixture components. The effects on the simple race–complex race competition of a cost of virulence, of density dependence and of differential adaptation were studied. The selection for simple or complex races in the pathogen population did not depend on initial race frequencies. For a given multiplication rate, complex race frequency increased faster when the spore dispersal gradient was shallow, when distribution of initial disease was generalized, when amount of initial disease was reduced and when the number of mixture components was increased. This was attributed to a better efficacy of the mixture in controlling simple races, resulting in a higher relative fitness of the complex race. For measured values of density dependence or differential adaptation effects, the complex race was at a higher frequency after a mean number of pathogen cycles between 2.5 and 5. The effect of the cost of virulence was stronger and, in certain situations, could result in selection for simple races. In the conditions of our simulations and with the effects tested, stabilization of the pathogen population in host mixtures was unlikely to occur. However, more information is needed concerning the rate at which complex races could evolve and how quickly mixture resistance could be eroded.     

Lesion-based analysis of leaf blast suppression in mixture of rice cultivar and a resistant near-isogenic line

Leaf blast suppression in multilines was evaluated based on the number of susceptible lesions observed in a pure stand of susceptible rice cultivar Sasanishiki, and in 1 : 1 and 1 : 3 mixtures of Sasanishiki and a resistant near-isogenic line, Sasanishiki BL4 or BL7, from 1998 to 2001. The number of lesions first observed in fields in the 1 : 1 and 1 : 3 mixtures were close to theoretical numbers calculated using the number of lesions observed in the pure stands and the ratios of the susceptible Sasanishiki in the mixtures. The ratio of the number of lesions in the 1 : 1 and 1 : 3 mixtures to the number in the pure stand was 0.29 and 0.09, respectively. The relationship between these ratios and the ratios of susceptible Sasanishiki in mixtures was defined in an equation to estimate the degree of leaf blast suppression. Validation studies for the ratios of the number of lesions in the 1 : 1 and 1 : 3 mixtures to the number in the pure stand were conducted in two different locations and showed that the ratios are almost acceptable. The calculated autoinfection to alloinfection ratio was 1.3 and 1.4 in the 1 : 1 and 1 : 3 mixtures, respectively, suggesting that the calculated ratio will affect the degree of leaf blast suppression. Thus, predictors were obtained to estimate leaf blast suppression for effective blast control in multilines.     

Synergism in fungicide mixtures againstPseudoperonospora Cubensis

Mixtures of cymoxanil with mancozeb (Mancur) or with mancozeb plus oxadixyl (Pulsan, Sandocur-M) were effective in controlling either metalaxyl-sensitive (MS) or -resistant (MR) isolates ofPseudoperonospora cubensis on cucumbers (cv. ‘Elem’) in growth chambers. Metalaxyl+mancozeb (Ridomil-MZ) and oxadixyl+mancozeb (SAN-518) mixtures were effective against the metalaxyl-sensitive but less so against the metalaxyl-resistant isolate. Fungicide mixtures were far more effective in controlling either the sensitive or the resistant isolates than were the individual components applied alone, thereby representing significant levels of synergistic interactions. Increased efficacy of mixtures relative to their combined components (synergy factor) was calculated by the Wadley method. Mixtures containing cymoxanil showed high synergy factors against the resistant isolate, whereas Ridomil-MZ and SAN-518 did not evince this phenomenon. We concluded that cymoxanil mixtures might be suitable for the control of downy mildew incited by a metalaxyl-resistant isolate ofP. cubensis.     

协同增效型拮抗细菌组合CL-7和CL-8的稳定性及其对加工番茄的促生防病效果

将加工番茄根际土壤中筛选到的单一拮抗细菌进行复配，筛选获得两个具有促生拈抗作用的细菌组合CL-7和CL-8。经过20代转管连续培养检测细菌组合的稳定性，结果表明，这两个拮抗细菌组合从13代之后，其pH值基本稳定，组合内菌落种类及数目也基本稳定；且12代后抑菌活性基本保持不变。和单一拮抗菌株相比，CL-7和CL-8均能显著增加对番茄立枯病菌、辣椒疫霉病菌和枯萎病菌的抑菌效果。发芽和苗期灌根试验结果表明，拮抗细菌组合CL-7和CL-8可显著促进加工番茄幼芽及苗期的生长，苗期鲜重分别增加了147．620k和176．190k，干重分别增加了143．75％和143．75％。在温室和田间条件下，细菌组合CL-7和CL-8对加工番茄立枯病的防治效果都很显著，分别可达25．1％、40．1％和52．2％、55．5％。     

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.     

Binary mixtures of pyrethroids produce differential effects on Ca influx and glutamate release at isolated presynaptic nerve terminals from rat brain

Isolated rat brain synaptosomes were used to evaluate the action of pyrethroid mixtures on Ca²⁺ influx and subsequent glutamate release under depolarizing conditions. In equipotent binary mixtures at their respective and/or estimated EC₅₀s with deltamethrin always as one of the two components, cismethrin, λ-cyhalothrin, cypermethrin, esfenvalerate and permethrin were additive and S-bioallethrin, fenpropathrin and tefluthrin were less-than-additive on Ca²⁺ influx. In binary mixtures with deltamethrin always as one of the two components, esfenvalerate, permethrin and tefluthrin were additive and λ-cyhalothrin was less-than-additive on glutamate release. Binary mixture of S-bioallethrin and cismethrin was additive for both Ca²⁺ influx and glutamate release. Only a subset of pyrethroids (S-bioallethrin, cismethrin, cypermethrin, and fenpropathrin) in binary mixtures with deltamethrin caused a more-than-additive effect on glutamate release. These binary mixtures were, however, only additive (cismethrin and cypermethrin) or less-than-additive (S-bioallethrin and fenpropathrin) on Ca²⁺ influx. Therefore, increased glutamate release evoked by this subset of pyrethroids in binary mixture with deltamethrin is not entirely occurring by Ca²⁺-dependent mechanisms via their action at voltage-sensitive calcium channels. These results suggest that pyrethroids do not share a common mode of toxicity at presynaptic nerve terminals from rat brain and appear to affect multiple target sites, including voltage-sensitive calcium, chloride and sodium channels.     

Resistance to acylalanines in Phytophthora infestans in The Netherlands1

In The Netherlands in 1980 a severe late-blight epidemic involving metalaxyl-resistant strains of Phytophthora infestans did considerable damage to the potato crop. As a consequence metalaxyl or metalaxyl-containing products were withdrawn from the Dutch fungicide market for the control of potato late blight. In 1981 the majority of the P. infestans isolates obtained from various parts of the country were sensitive to metalaxyl. In three areas where metalaxyl mixtures were used on a relatively large scale metalaxyl-resistant strains dominated the population. Incidental use of metalaxyl mixtures in 1982 and 1983 showed adequate late-blight control and, when in 1984 an early and in some areas severe epidemic developed, a metalaxyl/mancozeb mixture was reintroduced for curative application in combination with conventional fungicides. Although disease control was adequate, resistant strains were present in a small number of fields. Whether resistant strains will increase in frequency and threaten the usefulness of mixtures of acylalanines will heavily depend on how well potato farmers adopt strategies involving a limited use of mixtures of acylalanines in a spray schedule with conventional late-blight fungicides.     

Quantitative evolution of aggressiveness of powdery mildew under two-cultivar barley mixtures

The effects of cultivar mixtures on the evolution of aggressiveness of barley powdery mildew ( Blumeria graminis f.sp . hordei ) were modelled. It was found that the rate and direction of evolution of pathogen aggressiveness in a race-non-specific system and value at equilibrium, depends on initial resistance levels, proportions of component cultivars, autodeposition rates, the relative magnitude of the benefit of autoinfection, and the cost of alloinfection of spores. In the model, mixing cultivars in any proportions tended to reduce the aggressiveness of pathogens at equilibrium compared with pure stands, but this effect decreased when two mixture components were extremely unbalanced in proportion. Under low and medium autodeposition rates, the best control of the evolution of the pathogen was achieved by mixing two components in roughly equal proportions. The magnitude of aggressiveness at equilibrium increased as autodeposition rates increased. Though the level of initial resistance of mixture components did not have an impact on the aggressiveness of pathogens at equilibrium, it strongly influenced the transient values of aggressiveness and therefore the total amount of disease caused over an evolutionary period. The cost to the pathogen of alloinfection and benefit of autoinfection per se did not affect the final level of aggressiveness, but did affect the time to reach equilibrium. However, the ratio of the cost to the benefit did influence the final aggressiveness of the pathogen.     

Evaluation of the combined toxicity of 15 pesticides by uniform design

BACKGROUND: Environmental pesticides, including insecticides and herbicides, are frequently encountered as mixtures and threaten non‐target organisms in water. Evaluation of the combined toxicity of diverse pesticides with various concentration combinations is important, especially using limited experimental effort. Uniform design (UD) is one optimal experimental technique that can rationally arrange the concentrations of mixture components so that, with a minimum number of experimental runs, the combined toxicity of multiple pesticide mixtures can be evaluated. RESULTS: The concentration compositions of 18 pesticide mixture points designed by UD covered almost all possible concentration ranges of the mixture components on account of the two merits of ‘space filling’ and ‘multiple levels’. The combined toxicities of 18 mixture rays extended by using the fixed‐ratio ray design (FRRD) from 18 UD mixture points were evaluated by concentration addition (CA) and independent action (IA) models. It was found that the concentration–response curves (CRCs) predicted by CA were, on the whole, located between the 95% confidence intervals of the experimental CRCs, which implied that the combined toxicity of the pesticide mixture rays could be evaluated by CA. The CRCs predicted by IA were very similar to those from CA. CONCLUSION: The model developed from the UD mixture rays can effectively simulate mixtures with arbitrary concentration compositions of 15 pesticides. The CA model can accurately evaluate and predict the combined toxicity of the pesticides, which provides a useful tool for risk assessment of a mixture of multiple pesticides in the aquatic environment. Copyright © 2010 Society of Chemical Industry     

Effects of fungicide dose and mixtures on selection for triazole resistance in Mycosphaerella graminicola under field conditions

The effects of varying doses of fungicides, alone or in mixtures, on selection for triazole resistance were examined under field conditions. Two experiments were conducted using the triazole fungicide fluquinconazole with the strobilurin fungicide azoxystrobin as a mixture partner. Inoculated wheat plots with a known ratio of more sensitive to less sensitive isolates of the leaf blotch fungus Mycosphaerella graminicola were sprayed with fungicide and sampled once symptoms had appeared. Selection for fluquinconazole resistance increased in proportion to the dose, up to one-half of the full dose (the maximum tested) in both experiments. At the higher doses of fluquinconazole, the addition of azoxystrobin was associated with a decrease in selection (nonsignificant in the first experiment) for triazole resistance. Control by low doses of fluquinconazole was increased by mixture with azoxystrobin, but at higher doses mixture with azoxystrobin sometimes decreased control, so that reduced selection was obtained at the cost of some reduction in control. The effects on resistance are not necessarily general consequences of mixing fungicides, and suggest that the properties of any specific mixture may need to be demonstrated experimentally. Selection was inversely related to control in the unmixed treatments in both experiments, but the relationship was weaker in the mixtures with azoxystrobin.     

布氏白僵菌防治蔗田蛴螬的初步研究

采用甘蔗蛴螬残留虫口数和蔗根受害率的相对防效为指标,从蛴螬和甘蔗两个角度评价布氏白僵菌［Beauveria brongniartii (Sacc.) Petch］对甘蔗蛴螬的防治效果。结果表明,5月底6月初结合甘蔗大培土,布氏白僵菌粉剂单一施用及其与3.6%杀虫双颗粒剂混合施用,均可不同程度降低蔗田蛴螬虫量和蔗根受害率。布氏白僵菌粉剂单一施用时,以30.00 kg/hm2制剂用量的防效为最高,蔗田蛴螬残留虫口数和蔗根受害率的相对防效分别为67.27%和74.56%。两剂混合施用相对防效分别为74.55%～85.45%和78.18%～85.94%,其中以剂量为（22.50+45.00）kg/hm2的布氏白僵菌粉剂+3.6%杀虫双颗粒剂混合施用较为理想。     

Two‐component cultivar mixtures reduce rice blast epidemics in an upland agrosystem

The effect of two‐component rice cultivar mixtures on the control of rice blast disease was studied in three different experiments under rainfed upland conditions in the Madagascar Highlands. The mixtures involved a susceptible cultivar (either susceptible or very susceptible) and a resistant cultivar in different mixture arrangements (random or row mixtures) and with different proportions of the susceptible cultivar (50, 20 and 16·7%), which were compared to the susceptible cultivar grown in a pure stand. The effect of these mixtures on the incidence and severity of leaf and panicle blast was measured weekly, and on yield and yield components at harvest time. The mixture effect was more efficient in reducing disease with a proportion of 16·7% susceptible component than with a proportion of 50%. Blast epidemic was significantly reduced in all three experiments. However, under high blast pressure, there was no reduction in the disease by the end of the epidemic and yields of the susceptible cultivar were almost zero whatever the mixture. In two other experiments performed under lower blast pressure, disease incidence and severity were significantly lower in mixtures, and yields of the susceptible cultivars grown in mixtures were higher than those of their respective pure stands. Cultivar mixtures are a promising strategy that could contribute to a more sustainable cultivation of rice under upland conditions in the context of subsistence agriculture in Madagascar, where all cropping operations are manual.     

Selection of pathotypes of Erysiphe graminis f.sp. hordei in pure and mixed stands of spring barley

Pathotypes of Erysiphe graminis f.sp. hordei were monitored at fortnightly intervals in pure and mixed stands of spring barley during the course of mildew epidemics in two field trials. Mixtures were composed of cultivars with Arabische (gene Mla12 ), Laevigatum ( Ml(La) ), and Monte Cristo ( Mla9 ) resistance, respectively. The three-way mixtures were either random or, in 1989, laid out as one-row mixtures (i.e., regularly alternating rows of different genotypes) or three-row mixtures (i.e., regularly alternating three-row strips of different genotypes), respectively. In 1990 only random mixtures and six-row mixtures were compared with pure stands. The virulence complexity (i.e., the average number of virulence factors per isolate with reference to Mla12, Ml(La) , and Mla9 ) was always maximal in the random mixtures. In 1989, linear regression of complexity on mildew generations gave significant b -values (slopes) of 0·049, 0·031, and 0·025 in the random mixture, one-row mixture, and three-row mixture, respectively; the b -value from pure stands was not significant. In 1990, another sampling technique allowed selection to be observed on each genotype in the mixtures separately. In the random mixture b -values were 0·048, 0·064 and 0·017 (not significant) on Mla12, Ml(La) , and Mla9 cultivars respectively. In six-row mixtures and in pure stands, there was no significant increase in complexity ( b > 0) on any of the mixture components. Although the frequency and relative fitness of complex pathotypes were higher in all types of mixtures than in pure stands, selection towards complex races was much less intense in row mixtures than in random mixtures in both field trials.