Selection of dysfunctional alleles of bHLH1 and MYB1 has produced white grain in the tribe Triticeae

Grain color is a key agronomic trait that greatly determines food quality. The molecular and evolutionary mechanisms that underlie grain-color regulation are also important questions in evolutionary biology and crop breeding. Here, we confirm that both bHLH and MYB genes have played a critical role in the evolution of grain color in Triticeae. Blue grain is the ancestral trait in Triticeae, whereas white grain caused by bHLH or MYB dysfunctions is the derived trait. HvbHLH1 and HvMYB1 have been the targets of selection in barley, and dysfunctions caused by deletion(s), insertion(s), and/or point mutation(s) in the vast majority of Triticeae species are accompanied by a change from blue grain to white grain. Wheat with white grains exhibits high seed vigor under stress. Artificial co-expression of ThbHLH1 and ThMYB1 in the wheat endosperm or aleurone layer can generate purple grains with health benefits and blue grains for use in a new hybrid breeding technology, respectively. Our study thus reveals that white grain may be a favorable derived trait retained through natural or artificial selection in Triticeae and that the ancient blue-grain trait could be regained and reused in molecular breeding of modern wheat.