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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. [ABSTRACT FROM AUTHOR]

Mixing cultivars is an agroecological practice of crop diversification, increasingly used for cereals. The yield of such cereal mixtures is higher on average than the mean yield of their components in pure stands, but with a large variance. The drivers of this variance are plant-plant interactions leading to different plant phenotypes in pure and mixed stands, i.e phenotypic plasticity. The objectives were (i) to quantify the magnitude of phenotypic plasticity for yield in pure versus mixed stands, (ii) to identify the yield components that contribute the most to yield plasticity, and (iii) to link such plasticities to differences in functional traits, i.e. plant height and flowering earliness. A new experimental design based on a precision sowing allowed phenotyping each cultivar in mixture, at the level of individual plants, for above-ground traits throughout growth. Eight commercial cultivars of Triticum aestivum L. were grown in pure and mixed stands in field plots repeated for two years (2019–2020, 2020–2021) with contrasted climatic conditions and with nitrogen fertilization, fungicide and weed removal management strategies. Two quaternary mixtures were assembled with cultivars contrasted either for height or earliness. Compared to the average of cultivars in pure stands, the height mixture strongly underyielded over both years (-29%) while the earliness mixture overyielded the second year (+11%) and underyielded the first year (-8%). The second year, the magnitude of cultivar's grain weight plasticity, measured as the difference between pure and mixed stands, was significantly and positively associated with their relative yield differences in pure stands (R2=0.51). When grain weight plasticity, measured as the log ratio of pure over mixed stands, was partitioned as the sum of plasticities in each yield component, its strongest contributor was the plasticity in spike number per plant (∼56% of the sum), driven by even stronger but opposed underlying plasticities in both tiller emission and regression. For both years, the plasticity in tiller emission was significantly, positively associated with the height differentials between cultivars in mixture (R2=0.43 in 2019–2020 and 0.17 in 2020–2021). Plasticity in the early recognition of potential resource competitors is a major component of cultivar strategies in mixtures, as shown here for tillering dynamics. Our results also highlighted a link between plasticity in tiller emission and height differential in mixture. Both height and tillering dynamics displayed plasticities typical of the shade avoidance syndrome. Both the new experimental design and decomposition of plasticities developed in this study open avenues to better study plant-plant interactions in agronomically-realistic conditions. This study also contributed a unique, plant-level data set allowing the calibration of process-based plant models to explore the space of all possible mixtures. • A new field design allowed non-destructive phenotyping of individual plants in mixed stands. • Height and tillering were dynamically monitored in pure and mixed wheat stands. • Both over-yielding and under-yielding mixtures were observed over two mixtures and seasons. • The plasticity in spike number was the main contributor to such variations in yield. • Active plasticity due to the shade avoidance syndrome drives life-history strategies in mixture. [ABSTRACT FROM AUTHOR]

The reduction in biodiversity owing to agricultural intensification has brought negative effects on sustainable crop production in the agro-environment. Plant diversity can help regulate insect pests; however, research that demonstrates that genotypic diversity of multiple varieties with prominent agroecological practices to suppress insect pests and benefit crop yield at agricultural landscapes is limited. To examine the effects of diversity of wheat varieties on aphids and crop productivity, we performed field experiments by designing plots of single (resistant/susceptible) and cultivar mixtures in 2018–2020. The effects of population abundance of cereal aphids (Sitobion miscanthi and Rhopalosiphum padi) and crop yield on resistant and susceptible wheat varieties and mixture of wheat cultivars were determined to reveal the most dominant mixed broadcasting mode in Kaifeng of China. The results showed that cultivar mixture significantly decreased aphid abundance compared with the single varieties (resistant/susceptible). Cultivar mixture had a higher yield compared with the single varieties (resistant/susceptible). Additionally, cultivar mixture had significantly higher thousand-grain weights than susceptible and resistant wheat varieties in 2019 and 2020. Cultivar mixture enhanced wheat yield by decreasing cereal aphids, indicating a bottom-up or top-down effect from genetic diversity to pest abundance. Our results indicated that appropriate cultivar mixtures could manage insect pests to some extent and stabilize crop yield.

• Background and Aims Growing cultivars differing by their disease resistance level together (cultivar mixtures) can reduce the propagation of diseases. Although architectural characteristics of cultivars are little considered in mixture design, they could have an effect on disease, in particular through spore dispersal by rain splash, which occurs over short distances. The objective of this work was to assess the impact of plant height of wheat cultivars in mixtures on splash dispersal of Zymoseptoria tritici, which causes septoria tritici leaf blotch. • Methods We used a modelling approach involving an explicit description of canopy architecture and splash dispersal processes. The dispersal model computed raindrop interception by a virtual canopy as well as the production, transport and interception of splash droplets carrying inoculum. We designed 3-D virtual canopies composed of susceptible and resistant plants, according to field measurements at the flowering stage. In numerical experiments, we tested different heights of virtual cultivars making up binary mixtures to assess the influence of this architectural trait on dispersal patterns of spore-carrying droplets. • Key Results Inoculum interception decreased exponentially with the height relative to the main inoculum source (lower diseased leaves of susceptible plants), and little inoculum was intercepted further than 40 cm above the inoculum source. Consequently, tall plants intercepted less inoculum than smaller ones. Plants with twice the standard height intercepted 33 % less inoculum than standard height plants. In cases when the height of suscpeptible plants was doubled, inoculum interception by resistant leaves was 40 % higher. This physical barrier to spore-carrying droplet trajectories reduced inoculum interception by tall susceptible plants and was modulated by plant height differences between cultivars of a binary mixture. • Conclusions These results suggest that mixture effects on spore dispersal could be modulated by an adequate choice of architectural characteristics of cultivars. In particular, even small differences in plant height could reduce spore dispersal. [ABSTRACT FROM AUTHOR]

Increasing the biodiversity of cropped plants is a key leverage for agroecology, aiming to replace chemical inputs by ecological processes and regulations. Cultivar mixtures are a straightforward way to increase within-crop diversity, but they have so far been poorly used by farmers and they are not encouraged by advisory services. Based on the methodology developed by Kiær et al. (2009), we achieved a meta-analysis of cultivar mixtures in wheat. Among the 120 publications dedicated to wheat, we selected 32 studies to analyze various factors that may condition the success or failure of wheat mixtures by calculating overyielding, i.e. the difference in productivity of a variety mixture compared with the weighted mean of its component varieties in pure stand. The analysis highlighted a significant global overyielding of 3.5%, which reached 6.2% in condition of high disease pressures. Overyielding was not affected by seeding density or plot size. Under high disease pressure, overyielding increased by 3.2% point per added component variety. Overyielding was respectively 5.3% and 3.3% higher for mixtures heterogeneous in disease resistance or phenology than for homogeneous ones, and did not vary when considering height. Overyielding reached its highest values in the 1980s and 1990s, which reflects the predominance of disease-focused studies during this period. Our results confirm that cultivar mixtures are a potential way to increase yield relatively to pure varieties, especially under low pesticide cropping systems. Literature suggests that mixture practice is impeded by the lack of general rules that could help to mixing varieties. To design such rules it is needed to (1) achieve new experiments manipulating the heterogeneity in variety traits, (2) determine experimentally the ecological mechanisms underlying mixture performance and (3) develop new models allowing testing and analyzing these mechanisms. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The reduction in biodiversity owing to agricultural intensification has brought negative effects on sustainable crop production in the agro-environment. Plant diversity can help regulate insect pests; however, research that demonstrates that genotypic diversity of multiple varieties with prominent agroecological practices to suppress insect pests and benefit crop yield at agricultural landscapes is limited. To examine the effects of diversity of wheat varieties on aphids and crop productivity, we performed field experiments by designing plots of single (resistant/susceptible) and cultivar mixtures in 2018–2020. The effects of population abundance of cereal aphids (Sitobion miscanthi and Rhopalosiphum padi) and crop yield on resistant and susceptible wheat varieties and mixture of wheat cultivars were determined to reveal the most dominant mixed broadcasting mode in Kaifeng of China. The results showed that cultivar mixture significantly decreased aphid abundance compared with the single varieties (resistant/susceptible). Cultivar mixture had a higher yield compared with the single varieties (resistant/susceptible). Additionally, cultivar mixture had significantly higher thousand-grain weights than susceptible and resistant wheat varieties in 2019 and 2020. Cultivar mixture enhanced wheat yield by decreasing cereal aphids, indicating a bottom-up or top-down effect from genetic diversity to pest abundance. Our results indicated that appropriate cultivar mixtures could manage insect pests to some extent and stabilize crop yield. [ABSTRACT FROM AUTHOR]

Abstract: Plant ecology theory predicts that growing seed mixtures of varieties (variety mixtures) may increase grain yields compared to the average of component varieties in pure stands. Published results from field trials of cereal variety mixtures demonstrate, however, both positive and negative effects on grain yield. To investigate the prevalence and preconditions for positive mixing effects, reported grain yields of variety mixtures and pure variety stands were obtained from previously published variety trials, converted into relative mixing effects and combined using meta-analysis. Furthermore, available information on varieties, mixtures and growing conditions was used as independent variables in a series of meta-regressions. Twenty-six published studies, examining a total of 246 instances of variety mixtures of wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.), were identified as meeting the criteria for inclusion in the meta-analysis; on the other hand, nearly 200 studies were discarded. The accepted studies reported results on both winter and spring types of each crop species. Relative mixing effects ranged from −30% to 100% with an overall meta-estimate of at least 2.7% (p <0.001), reconfirming the potential of overall grain yield increase when growing varieties in mixtures. The mixing effect varied between crop types, with largest and significant effects for winter wheat and spring barley. The meta-regression demonstrated that mixing effect increased significantly with (1) diversity in reported grain yields, (2) diversity in disease resistance, and (3) diversity in weed suppressiveness, all among component varieties. Relative mixing effect was also found to increase significantly with the effective number of component varieties. The effects of the latter two differed significantly between crop types. All analyzed models had large unexplained variation between mixing effects, indicating that the variables retrievable from the published studies explained only a minority of the differences among mixtures and trials. [Copyright &y& Elsevier]

Wheat cultivars and cultivar mixtures were grown in commercial fields naturally infested with Cephalosporium gramanium. Thirteen treatments, including nine cultivars and four two-way cultivar mixtures, were replicated four times in 3×30 m plots in each of six experiments (two locations×three winter wheat seasons). Cephalosporium stripe was quantified by estimating the percentage of prematurely ripening heads (whiteheads) in each plot. There was little variation in percentage of whiteheads among club wheat entries, which were less susceptible than the common soft white winter wheats. Common wheat entries varied from highly susceptible to moderately resistant. One cultivar mixture significantly (P<0.01) increased the percentage of whiteheads as compared to the mean of its component cultivars grown in pure stand in four of the six experiments; mixtures never significantly decreased the percentage of whiteheads. All four cultivar mixtures significantly reduced lodging in the one experiment where there was sufficient eyespot (caused by Pseudocercosporella herpotrichoides) to cause lodging. Averaged over the six experiments, the four cultivar mixtures yielded 7.5, 6.9, 3.4, and −1.7% more than the means of their component cultivars grown in pure stand; the highest yielding club wheat and the highest yielding common wheat treatment were both cultivar mixtures. Cultivars varied in parameters of yield stability, with no obvious difference between susceptible and resistant cultivars. Cultivar mixtures demonstrated greater yield stability than did the individual cultivars. [Copyright &y& Elsevier]

The push for sustainable intensification requires development of alternative strategies for managing crop production that improve yields while maintaining or enhancing environmental services. We propose that intraspecific mixtures potentially exploit landscape ecosystem services to help suppress pests, impede disease spread, and ultimately improve yields. To test this hypothesis, we conducted a three-year field study of wheat cultivar mixtures compared to sole cultivar stands and evaluated pest control, yield, and economic benefits. Arthropod pests were not abundant in our experiment and predator abundance varied unpredictably across mixtures and sole cultivar stands. We found that mixtures suppressed foliar diseases at a level comparable to that of a fungicide application but had less impact on head diseases. Yield and economic return over variable costs were similar for mixtures and sole cultivar stands (3875 vs 3817 Mg ha−1 of wheat, $473 vs $462 ha−1, respectively), indicating that mixtures may provide similar yields and economic returns. In the three years of this project, there was no substantial benefit of using mixtures, but neither was there a penalty, indicating that planting mixtures can minimize risks should insect pests or disease outbreak. • Arthropod pests and predators were similar in cultivar mixtures or monocultures. • Cultivar mixture suppressed foliar disease compared to average sole cultivar stand with or without pesticide application. • Yield and economic return over variable costs were similar for mixtures and monocultures, but with less variability. [ABSTRACT FROM AUTHOR]

• Cultivar mixture was more infested by aphids than the least susceptible cultivar. • Intercrops tended to be less infested by aphids than wheat monoculture. • Combining genetic and species diversity did not better than each individual practice. • Wheat yields were reduced in intercropping, but not in cultivar mixtures. Increasing intrafield plant diversity has been shown to regulate pest populations. Mixing wheat cultivars and intercropping winter wheat and white clover are both promising agroecological practices. On field experiments over two growing seasons, we combined both practices and examined the impact on aphid populations and on wheat production. Results show that combining intra- and interspecific diversity did not outperform each practice individually in reducing aphid populations. Taken separately, intercropping tended to have lower aphid infestation, while it was intermediate in cultivar mixtures. Yearly variation in climatic conditions impacted wheat and clover development, as well as the appearance of aphid peaks. Wheat yields and grain nitrogen content were reduced in intercropping by 10% and 7%, respectively, but not in cultivar mixtures. Our findings suggest that intrafield diversification may regulate wheat aphids to some extent, but combining two diversification practices did not result in an attractive trade-off between pest regulation and wheat production in real farming conditions. [ABSTRACT FROM AUTHOR]