High productivity of wheat intercropped with maize is associated with plant architectural responses

Mixed cultivation of crops often results in increased production per unit land area, but the underlying mechanisms are poorly understood. Plants in intercrops grow differently from plants in single crops; however, no study has shown the association between plant plastic responses and the yield advantage. Here, we assessed the productivity of wheat-maize intercropping as compared to sole wheat and sole maize, and the associated differences in wheat shoot and leaf traits. In two field experiments, intercrop wheat and maize were both grown in alternating strips consisting of six rows of wheat and two rows of maize. The traits of wheat plants in border rows of the strips were compared to the traits of plants in the inner rows as well as those in sole wheat. Leaf development, chlorophyll concentration and azimuth, as well as the final leaf and ear sizes, tiller dynamics of wheat and yield components of both crops were determined. The relative densities of wheat and maize in the intercrop were 0.33 and 0.67, respectively, but the corresponding relative yields compared to the respective monocultures were 0.46 for wheat and 0.77 for maize. Compared to wheat plants in the inner rows of the intercrop strips as well as in the monoculture, border-row wheat plants in the intercrop strips had (a) more tillers owing to increased tiller production and survival, and thus more ears, (b) larger top leaves on the main stem and tillers, (c) higher chlorophyll concentration in leaves, (d) greater number of kernels per ear and (e) smaller thousand-grain weight. Grain yield per metre row length of border-row wheat was 141% higher than the sole wheat, and was 176% higher than the inner-row wheat. The results demonstrate the importance of plasticity in architectural traits for yield advantage in multispecies cropping systems.