Contrasted reaction norms of wheat yield in pure vs mixed stands explained by tillering plasticities and shade avoidance

Abstract • Context 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. • Objectives 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 most to yield plasticity, and (iii) to link such plasticities to differences in functional traits, height and earliness. • Methods 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. • Results 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 slightly underyielded the first year (-8%). The second year, the magnitude of plasticity in grain weight was significantly and positively associated with their relative yield differences in pure stands (R 2 =0.51). The plasticity in spike number was the strongest contributor to yield plasticity (∼56%), driven by even stronger but opposed 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 (R 2 =0.43 in 2019-2020 and 0.17 in 2020-2021). • Conclusions Plasticity in the early recognition of potential competitors for resources is a major component of genotype 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. • Implications 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.