Effects of Wheat Cultivar Mixtures on Epidemic Progression of Septoria Tritici Blotch and Pathogenicity of Mycosphaerella graminicola

ABSTRACT The effects of host genotype mixtures on disease progression and pathogen evolution are not well understood in pathosystems that vary quantitatively for resistance and pathogenicity. We used four mixtures of moderately resistant and susceptible winter wheat cultivars naturally inoculated with Mycosphaerella graminicola to investigate impacts on disease progression in the field, and effects on pathogenicity as assayed by testing isolate populations sampled from the field on greenhousegrown seedlings. Over 3 years, there was a correspondence between the mixtures' disease response and the pathogenicity of isolates sampled from them. In 1998, with a severe epidemic, mixtures were 9.4% less diseased than were their component pure stands (P = 0.0045), and pathogen populations from mixtures caused 27% less disease (P = 0.085) in greenhouse assays than did populations from component pure stands. In 1999, the epidemic was mild, mixtures did not reduce disease severity (P = 0.39), and pathogen populations from mixtures and pure stands did not differ in pathogenicity (P = 0.42). In 2000, epidemic intensity was intermediate, mixture plots were 15.2% more diseased than the mean of component pure stands (P = 0.053), and populations from two of four mixtures were 152 and 156% more pathogenic than the mean of populations from component pure stands (P = 0.043 and 0.059, respectively). Mixture yields were on average 2.4 and 6.2% higher than mean component pure-stand yields in 1999 and 2000, respectively, but the differences were not statistically significant. The ability of mixtures challenged with M. graminicola to suppress disease appears to be inconsistent. In this system, host genotype mixtures evidently do not consistently confer either fitness benefits or liabilities on pathogen populations.     

Disease severity and yield of pure-line wheat cultivars and mixtures in the presence of eyespot, yellow rust, and their combination

Five winter wheat cultivars, six two-component cultivar mixtures, and one four-way mixture were grown in the presence of yellow rust, eyespot, both diseases, and neither disease for three seasons. On average, mixtures reduced the severity of yellow rust relative to their component pure stands by 53%. The four-component mixture provided better yellow rust control than did the two-way mixtures. Eyespot severity was reduced through mixing only in the absence of yellow rust and by only three of the seven mixtures (mean reduction = 13%). Yellow rust was 13% less severe in the presence of eyespot, and eyespot was 10% more severe in the presence of yellow rust. Averaged over all years, the mixtures increased yield relative to the pure stands by 6·2, 1·7, 7·1, and 1·3% in the presence of yellow rust, eyespot, both diseases, and neither disease, respectively. Two mixtures provided significant yield increases over the means of their component pure stands (7% and 9%) in the presence of eyespot even though one of them did not significantly reduce eyespot severity. Accounting for all disease treatments and years, four mixtures provided distinctly higher yield increases than the other three. In mixtures containing a resistant cultivar and a cultivar susceptible to eyespot, yield loss by the susceptible cultivar was not compensated for by increased yield of the resistant cultivar. The mixtures showed improved yield stability relative to the pure stands, with the four-component mixture being particularly stable.     

Cultivar mixture effects on disease and yield remain despite diversity in wheat height and earliness

Cultivar mixtures can stabilize yield and reduce pathogen spread in plant populations. A field experiment was performed to determine (a) whether a large difference between the cultivars in the mixture (e.g., plant height or earliness) would have an impact on mixture performance, and (b) whether such differences would modify the classical rules for mixture design. Mixtures were constituted from wheat cultivars with diversity for many traits, including plant height, flowering date, disease resistance, and yield potential. The field experiment was conducted over 3 years, testing each year 72–90 mixtures of two, four, or eight cultivars, and their corresponding pure stands. Disease severity and yield of cultivar mixtures were strongly related to the mean values of the component cultivars in pure stands. Despite the considerable diversity of the mixtures tested, the classic rules (e.g., proportion of susceptible cultivars) already tested in mixtures with similar height and earliness were effective for decreasing disease severity. Agronomic heterogeneity for traits such as plant height, yield potential, or earliness of the cultivars in mixtures did not have a negative impact on disease severity and yield relative to pure stands. Increasing the number of cultivars in the mixture from two to eight had no impact on the mean disease severity and yield of the mixtures, but reduced the variability of disease severity and yield in the mixture relative to pure stands. These results suggest that it may be possible to increase within-field wheat diversity by combining more contrasting cultivars in mixtures than was previously thought.     

Host frequency and density effects on powdery mildew and yield in mixtures of barley cultivars

The effects of frequency and density of susceptible plants on barley powdery mildew epidemics were studied in a combined set of addition and replacement series of field trials. In the addition series, plant densities in pure stands of three cultivars, Rambo, Rodos and Grosso (susceptible, moderately resistant and immune, respectively) were varied six-fold. In the replacement series, the three possible two-way mixtures were analysed at different frequencies but at a density corresponding to the maximum pure stand density. Disease and yield were assessed on a per-plant basis. In the pure stands, tillering reduced the range of densities from six-fold to between three- and four-fold, while in the mixtures, frequencies changed only slightly over time, indicating that competitive interactions among the cultivars were roughly equal. Yield per plant decreased logarithmically with increasing density as expected. However, yield per seed head was not correlated with the final number of heads per plot, indicating low competition among heads even at the highest density. Disease in susceptible pure stands increased strongly with decreasing density in 1994 and to a lesser degree in 1995. These differences could have been caused by differences in plant nutritional status and consequent epidemiological effects. Disease reduction on the susceptible cultivars in mixtures varied between 33% and 71% among years. Depending on the length and strength of the epidemic, the effects of host density and frequency on disease severity varied substantially among years.     

Effect of two-component cultivar mixtures and yellow rust on yield and yield components of wheat

Four winter wheat ( Triticum aestivum ) cultivars and three two-component cultivar mixtures were planted in a replacement series both inoculated with or protected from yellow rust ( Puccinia striiformis ) in three environments. Each cultivar was susceptible to one or two of the rust races used. Mixtures yielded, on average, 7 and 4% more than their component pure stand means under inoculated and rust-free conditions, respectively. Though all yield components were affected by yellow rust, seed weight was the component that was most consistently influenced. The component genotypes within mixtures varied considerably with respect to yield, and the yield of the same component cultivar included in different mixtures sometimes differed significantly. The correlation between yellow rust severity/tiller and grain yield/tiller in mixture differed among cultivars and depended on their companion cultivar. Variance component analysis indicated that yellow rust was the most important experimental variable influencing grain yield. There was no relationship between yield of the cultivars in pure stands and their yields or competitive abilities in mixture. Disease did not change the competitive ranking of cultivars in mixture. Mixtures with complementary, negative, and overcompensatory interactions were identified. On average, mixtures showed no greater yield stability than did pure stands.     

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.     

Incidence of Soil-borne Wheat Mosaic Virus in Mixtures of Susceptible and Resistant Wheat Cultivars

The multiplication of Soil-borne wheat mosaic virus (SBWMV) was studied in mixtures of two winter wheat (Triticum aestivum) cultivars, one susceptible (Soissons) and the other resistant (Trémie). Two seed mixtures of susceptible and resistant varieties in ratios of 1 : 1 and 1 : 3 and their component pure stands, i.e. each variety grown separately, were grown in a field infected with SBWMV. The presence of the virus was detected using DAS-ELISA from January to May. The resistant cultivar Trémie showed no foliar symptoms nor could the virus be detected in the leaves or roots. In May, about 88% of plants of susceptible cultivar Soissons grown in pure stands were infected. At this time, the disease reduction relative to pure stands was 32.2% in the 1 : 1 mixture and 39.8% in the 1 : 3 mixture. Optical density (OD) values from ELISA of the infected plants in the two mixtures were consistently lower than that of the infected plants in cultivar Soissons in pure stands. The ELISA index (EI) calculated using three scales of OD values was 65.5% in the susceptible cultivar in pure stands. The value for this index was 19.1% in the 1 : 1 mixture and 7.9% in the 1 : 3 mixture. The plants of the resistant cultivar Trémie infected in the same field and transferred in January to a growth cabinet at 15 °C multiplied the virus and produced viruliferous zoospores. These results show that the resistant cultivar Trémie plays a role in disease reduction in the cultivar mixtures in field conditions. Possible reasons for this are discussed.     

Protection of susceptible potato cultivars against late blight in mixtures increases with decreasing disease pressure

ABSTRACT Cultivar mixtures can reduce potato late blight severity on susceptible cultivars. While alternating rows of susceptible and resistant cultivars would be more acceptable than random mixtures for commercial use, they increase the genotype unit area, which is an unfavorable factor for mixture efficiency, and have been minimally efficient when disease pressure is high. The effects of disease pressure on the performance of alternating rows of cultivars possessing various types and levels of resistance were investigated in 2000 and 2001 near Quito, Ecuador, where natural pressure of late blight is high. The experiments included the highly susceptible cvs. Cecilia in 2000 and LBr37 in 2001, as well as C114 (moderately resistant) and PAN (highly resistant), planted as pure stands and as the three possible two-way combinations. Different disease pressures were obtained with three spraying schedules of a contact fungicide: nontreated, one spray every second week, and one spray weekly. The area under the disease progress curve (AUDPC) on the susceptible cultivar was 0 to 20% less in mixed than in pure plots when no fungicide was applied, 13 to 26% less with a biweekly application of fungicide, and 32 to 53% less with a weekly application. These values are comparable to those obtained in previous experiments in smaller plots with designs maximizing the distance between susceptible plants. No significant differences in mixture performance were observed according to the resistant cultivar included. Effects on yield were minimal, because of the impact of factors other than late blight. Disease pressure therefore appears as a major factor conditioning the efficiency of potato cultivar mixtures against late blight.     

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.     

Spatiotemporal effects of cultivar mixtures on wheat stripe rust epidemics

The use of cultivar mixtures is increasingly practical in wheat stripe rust management. Field experiments with wheat cultivar mixtures were conducted to determine their effects on temporal and spatial patterns of stripe rust epidemics in three regions. In the Beijing and Gangu fields, where the epidemics were caused by artificial inoculation, disease incidence and the area under the disease progress curve (AUDPC) of the cultivar mixtures were significantly lower (P < 0.05) than those of the susceptible pure stands. We defined the relative effectiveness of cultivar mixture on disease development related to that in pure stands (REM). The results demonstrated that in many treatments of mixtures of susceptible cultivar with resistant cultivars at various ratios in different locations, their effects on disease reduction were positive (REM < 1). The reduction of epidemic rate in cultivar mixtures expressed in either early season or late season depended on the initial pattern of disease and cultivar mixture treatments. Semivariograms were used to determine the spatiotemporal patterns of disease in the Gangu field. The spatial analysis showed clear spatial patterns of the disease in all four directions of the fields on susceptible pure stands but not on cultivar mixtures. The results implied that the mechanisms of cultivar mixture on disease management might include the interruption of disease spatial expansion and a physical barrier to pathogen inoculum by resistant cultivars.     

Panicle blast and canopy moisture in rice cultivar mixtures

ABSTRACT Glutinous rice cultivars were sown after every fourth row of a nonglutinous, hybrid cultivar in an additive design. The glutinous cultivars were 35 to 40 cm taller and substantially more susceptible to blast than was the nonglutinous cultivar. Interplanting of glutinous and nonglutinous rice reduced the incidence and severity of panicle blast on the glutinous cultivars by >90%, and on the nonglutinous cultivar by 30 to 40%. Mixing increased the per unit area yield of glutinous rice by 80 to 90% relative to pure stand, whereas yield of the nonglutinous cultivar was essentially unaffected by mixing. To determine whether the different plant heights and canopy structures may contribute to a microclimate that is less favorable to blast infection, we monitored the moisture status of the glutinous cultivars in pure stand and mixture at 0800 h by measuring relative humidity at the height of the glutinous panicles using a swing psychrometer and by visually estimating the percentage of leaf area covered by dew. Averaged over the two seasons, the number of days of 100% humidity at 0800 h was 20.0 and 2.2 for pure stands and mixtures, respectively. The mean percentage of glutinous leaf area covered by dewwas 84 and 36% for the pure stands and mixtures, respectively. Although other mechanisms also were operative, reduced leaf wetness was likely a substantial contributor to panicle blast control in the mixtures.     

Effects of disease and plant competition on yield in monocultures and mixtures of two wheat cultivars

Disease seventy and crop yield were compared in monocultures and mixtures of two wheat cultivars. one susceptible and one resistant to Puccinia graminis f. sp. tritici. Part of each plot was inoculated and part treated with fungicide. Rust seventy was inversely related to the proportion of resistant plants present. Weight of grain in 100 heads of the susceptible cultivar increased as the amount of rust decreased. The susceptible cultivar yielded less than the resistant cultivar in monoculture, but was a stronger competitor in mixtures. Competitive abilities of the cultivars were independent of severity of rust in the mixtures studied.     

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

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

Improved control of septoria tritici blotch in durum wheat using cultivar mixtures

Mixtures of cultivars with contrasting levels of resistance can suppress infectious diseases in wheat, as demonstrated in numerous field experiments. Most studies focus on airborne pathogens in bread wheat, while splash-dispersed pathogens have received less attention, and no studies have been conducted in durum wheat. We conducted a 2-year field experiment in Tunisia to evaluate the performance of cultivar mixtures with varying proportions of resistance (0%–100%) in controlling the polycyclic, splash-dispersed disease septoria tritici blotch (STB) in durum wheat. To measure STB severity, we used a high-throughput method based on digital image analysis of 3,074 infected leaves collected from 42 and 40 experimental plots during the first and second years, respectively. This allowed us to quantify pathogen reproduction on wheat leaves and to acquire a large data set that exceeds previous studies with respect to accuracy and precision. Our analyses show that introducing only 25% of a disease-resistant cultivar into a pure stand of a susceptible cultivar provides a substantial reduction of almost 50% in disease severity compared to the susceptible pure stand. However, incorporating two resistant cultivars instead of one did not further improve disease control, contrary to predictions of epidemiological theory. Susceptible cultivars can be agronomically superior to resistant cultivars or be better accepted by growers for other reasons. Hence, if mixtures with only a moderate proportion of the resistant cultivar provide a similar degree of disease control as resistant pure stands, as our analysis indicates, such mixtures are more likely to be accepted by growers.     

Effect of two-component cultivar mixtures on development of wheat yellow rust disease in the field and greenhouse in the Nepal Himalayas

Wheat yellow rust (WYR), caused by Puccinia striiformis f. sp. tritici (PST), is a major disease of wheat, and deployment of a single cultivar often leads to disease epidemics. Effect of inoculum level, foliar fungicide spray, and wheat cultivar mixtures were evaluated on disease development in the field and greenhouse in Nepal. Treatments were arranged in a split–split plot design with three replications in both experiments. Two inoculum levels of PST (low and high) were main plot factors; nontreated control and foliar spray of fungicides (Mancozeb and Bayleton) were subplot factors; and two-component cultivar mixtures, composed of different ratios of a susceptible (S) and a resistant (R) cultivars (90:10, 80:20, and 50:50, 100:0, and 0:100) were sub–subplot factors. WYR severity was assessed at different time intervals, and disease development was calculated as area under the disease progress curve (AUDPC). Inoculum level did not cause significant differences in AUDPC in the field but did in the greenhouse. Foliar spray of fungicides reduced the AUDPC in the greenhouse and field. In both experiments, AUDPC values were low in cultivar mixtures compared with a pure stand of a susceptible cultivar. As the proportion of resistant cultivar increased compared with the susceptible cultivar in the S:R mixture component, disease severity decreased with a consequent increase in grain yield. The greater yield obtained with cultivar mixtures compared with only the susceptible cultivar, independent of inoculum level and fungicide spray in the field, revealed a promising strategy to manage WYR in Nepal.     

Epidemiological effect of gene deployment strategies on bacterial blight of rice

ABSTRACT Experiments were conducted in farmers' fields at two locations of the irrigated lowlands of Laguna province in southern Luzon island, Philippines, during the wet seasons of 1993 and 1994. Nine rice populations were studied including pure stands, two-component mixtures, two-gene combinations of backcrossed lines containing varying combinations of the bacterial blight resistance genes Xa-4, xa-5, and Xa-10, and a non-isogenic cultivar containing Xa-4 and partial resistance to bacterial blight. The area under the disease progress curve (AUDPC) of both gene combinations studied was significantly less than the single most effective gene of each combination deployed singly. A mixture of a susceptible and a resistant line expressed an AUDPC significantly less than the mean of its component pure stands, but two other mixtures did not. The cultivar IR20, which contains both Xa-4 and partial resistance, reduced the AUDPC by about two-thirds as compared with IR-BB4, which contains Xa-4 and little or no partial resistance.     

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.     

Effects of induced resistance on infection efficiency and sporulation of Puccinia striiformis on seedlings in varietal mixtures and on field epidemics in pure stands

An inducer race ofPuccinia striiformis inoculated two days before a challenger race on wheat seedlings of cv. Clement, reduced infection efficiency by 44% and lesion expansion by 7.7%; there was no effect on sporulation rate. In field epidemics, induced resistance restricted the disease intensity on cvs. Clement and Austerlitz in pure stands by 44% and 57% respectively. In the glasshouse at the seedling stage, disease intensity in a varietal mixture was reduced by 82% compared to the susceptible pure stands; one third of the total disease reduction was attributable to induced resistance.     

Effects of blast on components of wheat physiology and grain yield as influenced by fungicide treatment and host resistance

Two field experiments (Exp. 1 and Exp. 2) were carried out to assess the physiological performance and grain yield of wheat cultivars BR‐18 (moderately resistant) and Guamirim (susceptible) inoculated with Pyricularia oryzae in plots treated or untreated with Ópera (fungicide 13.3% epoxiconazole + 5% pyraclostrobin). Results from regression analyses indicated that spike and leaf blast severity at 10–14 days after inoculation (dai) were associated with greater yield losses (highest negative slope) than severity at 18–22 dai. Relative to untreated Guamirim, there were 0.3% and 16% increases in Exp. 1 and 2, respectively, for untreated BR‐18 (resistance alone). For fungicide treatment alone, the mean yield of Guamirim increased by 20% and 61% in Exp. 1 and 2, respectively, relative to the untreated fungicide control, whereas for the fungicide treated BR‐18, the mean yield increased by 26% and 83% in Exp. 1 and 2, respectively. Fungicide application and cultivar resistance resulted in higher measures of leaf health and photosynthetic performance in both spikes and leaves than in the untreated susceptible reference treatment. The results from this study may be useful in future efforts to develop crop loss models and management guidelines for wheat blast.