The development of hooded awns in barley: From ectopic Kap1 expression to yield potential

Awns in barley have different shapes including awnless, straight, hooded, crooked, and leafy awns. The hooded awns are characterized by an appendage of the lemma, which forms a trigonal or cap-shaped structure, and even blossoms and yields fruits on barley awn. In the lemma primordia of wild-type (straight awn), cells divide and elongate to form the straight awn. However, in the lemma primordia of KNOX3 mutant (hooded awn), cells divide at various orientations without elongating, and they form hooded awns. This phenomenon is due to the upregulation of KNOX3 expression via insertion of a tandem direct duplication of 305 bp in the intron IV. Here, we summarize the development of barley hooded awn research in the following two aspects: on the one hand, the morphology, development of hooded awns, and the expression regulation of the KNOX3 gene. The latter includes ectopic expression of the KNOX3 gene, gene interactions among awn-related genes, the regulatory relationship between class I KNOX genes and hormones, as well as the influence of abiotic stresses. On the other hand, the potential performance of hooded awns in barley for yield breeding is discussed. Hooded awns have potential application value in forage, which could compensate for the disadvantage of the long straight awn in the barley straw used for feed in modern cultivars. In addition, the hooded awn produces ectopic meristems to develop complete florets, which is an interesting question and helps to understand the development, adaptation, and evolution of plant floral organs.