Symmetric response to competition in binary mixtures of cultivars associates with genetic gain in wheat yield

The evolution in the definition of crop yield-from the ratio of seed harvested to seed sown to the contemporary measure of mass of seed per unit land area-has favoured less competitive phenotypes. Here we use binary mixtures of cultivars spanning five decades of selection for yield and agronomic adaptation to ask three questions. First, what is the degree of symmetry in the response of yield to neighbour; this is, if an older, more competitive cultivar increases yield by 10% with a less competitive neighbour in comparison to pure stands, would the newer, less competitive cultivar reduce yield by 10% when grown with older neighbour. Lack of symmetry would indicate factors other than competitive ability underly yield improvement. Second, what are the yield components underlying competitive interactions. Third, to what extent are the responses to neighbour mediated by radiation, water and nitrogen. A focus on yield components and resources can help the interpretation of shifts in the crop phenotype in response to selection for yield. The rate of genetic gain in yield over five decades was 24 kg ha^-1 year^-1 or 0.61% year^-1. A strongly symmetrical yield response to neighbour indicates that yield improvement closely associates with a reduction in competitive ability. Response to neighbour was larger for grain number and biomass than for grain weight and allocation of biomass to grain. Under our experimental conditions, competition for radiation was dominant compared to competition of water and nitrogen. High-yielding phenotypes had lower competitive ability for radiation but compensated with higher radiation use efficiency, a measure of canopy photosynthetic efficiency. Genetic and agronomic manipulation of the crop phenotype to reduce competitive ability could further improve wheat yield to meet the challenge of global food security.