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.     

Influence of barley variety mixtures on severity of scald and net blotch and on yield *

Two susceptible barley cultivars and two moderately resistant advanced breeding lines were grown as pure stands and as the 11 possible equi-proportional mixtures over three field seasons, in the presence of scald and net blotch. Plots were either inoculated with infested straw, or non-inoculated and sprayed with fungicide. On average, the variety mixtures restricted leaf disease development by 12%. However, mixtures of either or both susceptible cultivar/s with the moderately resistant line 1861018 restricted disease severity by 20-32%. Generally, the mixtures did not increase yield relative to the mean of the pure stands. However, in the year of highest disease severity, yield was increased by 7% owing to mixing in the inoculated plots. In addition, one mixture consistently increased yield (mean of 4%) and one mixture had consistently lower yields (mean of 6%)) over years and inoculation treatments. Thus, careful evaluation is required in this environment and with these genotypes in order to identify mixtures with positive effects on disease control and yield.     

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.     

小麦品种多样性对白粉病及产量和蛋白质的影响

 对品种多样性控制小麦白粉病的效果以及对产量和蛋白质的影响进行了研究,为利用品种多样性控制白粉病提供理论依据和技术支持。采用SSR技术分析5个小麦品种的亲缘关系;2008年和2009年按组合所含品种数为2、3、4和5个进行组合,在田间种植混种组合和单播小区,人工接种白粉菌,比较各小区中小麦白粉病的AUDPC值、产量和蛋白质变化。结果表明,5个品种亲缘关系相对较近;2个品种混合时以抗感搭配较好,随着抗病品种比例增加控病效果增强;2008年混合种植中有防治效果的组合占54.55%,相对防效为10.02%~47.58%,2009年有防治效果的组合占23.08%,相对防效在1.85% 到18.96 % 之间;在适当的组合中,如京冬8号/轮选987、京双16/京411/京冬8号对小麦白粉病有控制效果,混种不会影响产量和蛋白质含量;当有大量能克服本地品种抗性的外来菌源时混合种植的控病效果降低。品种多样性可以用做防治小麦白粉病的一项措施。     

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.     

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.     

Cultivar mixtures for the simultaneous management of multiple diseases: tan spot and leaf rust of wheat

ABSTRACT Because of differences in life histories between Puccinia triticina, a highly specialized, polycyclic, windborne pathogen with a shallow dispersal gradient, and Pyrenophora tritici-repentis, a residue-borne pathogen with a steep dispersal gradient, wheat mixtures are expected to be more effective at controlling leaf rust than tan spot. The objectives of this research were to determine the effect of two-cultivar mixtures with varying proportions and different pathogen resistance profiles on the severity of tan spot and leaf rust, to evaluate yield of the mixtures in the presence or absence of disease, and to directly compare the relative effectiveness of cultivar mixing for tan spot versus leaf rust. In a field experiment at two sites in Kansas over two growing seasons, winter wheat cvs. Jagger and 2145, which have differential resistance reactions to leaf rust and tan spot, each were planted in proportions of 0.25, 0.50, 0.75, and 1.00. Plots were inoculated with each pathogen alone, both pathogens, treated with a fungicide, or exposed to ambient conditions. For both diseases for all siteyears, severity decreased substantially on the susceptible cultivar as the proportion of that cultivar decreased in mixture. Mixtures were significantly more effective at reducing leaf rust than tan spot in three of four site-years. Mixtures generally yielded the same as the weighted mean of components in monoculture although, in two of three site-years, at least one fungicide-treated and one diseased mixture each yielded higher than expected values. Although this particular mixture produced only modest yield benefits, the potential for simultaneous reductions in tan spot and leaf rust was demonstrated.     

小麦品种混（间）种对小麦条锈病防治效果和产量影响研究

利用甘肃甘谷县和四川盐亭县当地各4个对条锈病具不同抗性的小麦品种,按照抗感品种不同比例混（间）种研究各种植方式对小麦条锈病的防治作用和对小麦的增产效果,并以REM比较各混（间）种种植方式的效果。结果表明,混（间）种对本地菌源引起的病害有明显效果,混种对病害的减轻程度为29.6％～81.9%,对外来菌源引起的病害效果较差;混（间）种对于小麦的增产作用也表现不一,28个混间种处理中,仅8个处理（占28.6%）有增产效果。     

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.     

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.     

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.     

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.     

小麦品种混种对白粉病的调控及其对部分产量性状及蛋白质含量的影响

对品种混合种植控制小麦白粉病的效果以及对小麦穗重、千粒重、蛋白质等指标的影响进行了研究,为混合种植控制白粉病提供理论依据和品种组合。在对37个小麦品种进行SSR多样性分析的基础上,选用8个生产品种（后备品种）,按品种数3、4、5、6、8进行组合,在田间以随机区组设计种植混播组合和单播小区,人工接种白粉菌,比较各小区中小麦白粉病的AUDPC值、穗重、千粒重和粗蛋白差异。结果表明,8个品种亲缘关系较远;4品种混种时AUDPC最小,穗重最重。26个品种混种组合中有防治效果的组合占到73.08%,相对防效为1.23%~56.65%;混种未对穗重和粗蛋白含量造成负面影响。品种多样性种植可以用做调控小麦白粉病的一项措施。     

Wheat leaf rust severity as affected by plant density and species proportion in simple communities of wheat and wild oats

ABSTRACT While it is generally accepted that dense stands of plants exacerbate epidemics caused by foliar pathogens, there is little experimental evidence to support this view. We grew model plant communities consisting of wheat and wild oats at different densities and proportions and exposed these communities to Puccinia recondita to induce wheat leaf rust. Wild oats was included because it is a common competitor of wheat and may act as a barrier to the dispersal of P. recondita spores among wheat plants. Disease severity was estimated as percentage of wheat flag leaves covered by rust lesions. Seeding density rarely had a significant influence on rust severity, probably because of compensation due to increased tillering at low seeding densities. In contrast, increasing the proportion of wheat in mixtures with wild oats consistently increased wheat leaf rust severity. Regression parameters describing wheat leaf rust severity as a function of wheat seeding density did not differ significantly between pure wheat stands and wheat-wild oat mixtures and, thus, failed to support an effect of wild oats on wheat leaf rust other than through its competitive impact on wheat tiller density.     

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.     

The impact of wheat cultivar mixtures on virulence dynamics in Zymoseptoria tritici populations persists after interseason sexual reproduction

This study follows on from a previous study showing that binary mixtures of wheat cultivars affect the evolution of Zymoseptoria tritici populations within a field epidemic from the beginning (t1) to the end (t2) of a growing season. Here, we focused on the impact of interseason sexual reproduction on this evolution. We studied mixtures of susceptible and resistant cultivars (carrying Stb16q, a recently broken-down resistance gene) in proportions of 0.25, 0.5 and 0.75, and their pure stands. We determined the virulence status of 1440 ascospore-derived strains collected from each cultivar residue by phenotyping on seedlings. Virulence frequencies in the ascospore-derived population were lower in mixtures than in pure stands of the resistant cultivar, especially in the susceptible cultivar residues, as at t2, revealing that the impact of mixtures persisted until the next epidemic season (t3). Surprisingly, after sexual reproduction the avirulence frequencies on the resistant cultivar residues increased in mixtures where the proportion of the susceptible cultivar was higher. Our findings highlight two epidemiological processes: selection within the pathogen population between t1 and t2 driven by asexual cross-contamination between cultivars (previous study) and sexual crosses between avirulent and virulent strains between t2 and t3 driven by changes in the probabilities of physical encounters (this study). Mixtures therefore appear to be a promising strategy for the deployment of qualitative resistances, not only to limit the intensity of Septoria tritici blotch epidemics, but also to reduce the erosion of resistances by managing evolution of the pathogen population at a pluriannual scale.     

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.