Preferential helping to relatives: A potential mechanism responsible for lower yield of crop variety mixtures?

Variety mixtures, the cultivation of different genotypes within a field, have been proposed as a way to increase within-crop diversity, allowing the development of more sustainable agricultural systems with reduced environmental costs. Although mixtures have often been shown to over-yield the average of component varieties in pure stands, decreased yields in mixtures have also been documented. Kin selection may explain such pattern, whenever plants direct helping behaviors preferentially toward relatives and thus experience stronger competition when grown with less related neighbors, lowering seed production of mixtures. Using varieties of durum wheat originating from traditional Moroccan agrosystems, we designed a greenhouse experiment to address whether plants reduced competition for light by limiting stem elongation when growing with kin and whether such phenotypic response resulted in higher yield of kin groups. Seeds were sown in groups of siblings and nonkin, each group containing a focal plant surrounded by four neighbors. At the group level, mean plant height and yield did not depend upon relatedness among competing plants. At the individual level, plant height was not affected by genetic relatedness to neighbors, after accounting for direct genetic effects that might induce among-genotype differences in the ability to capture resources that do not depend on relatedness. Moreover, in contrast to our predictions, shorter plants had lower inclusive fitness. Phenotypic plasticity in height was very limited in response to neighbor genotypes. This suggests that human selection in crops may have attenuated shade-avoidance responses to competition for light. Future research on preferential helping to relatives in crops might thus target social traits that drive competition for other resources than light. Overall, our study illustrates the relevance of tackling agricultural issues from an evolutionary standpoint and calls for extending such approaches to a larger set of crop species.