Genetic analysis of mutations that alter cell fates in maize leaves: dominant Liguleless mutations

The three major components of the maize leaf are the blade, the sheath, and at their junction, ligular region. Each exhibits specific cell types and organization. Four dominant Liguleless (Lg) mutations (Lg3-O, Lg4-O, Lg*347, and Lg*9167) in at least three different genes cause a similar morphological phenotype in leaves, although each mutation affects a distinct domain of the blade. Mutant leaves display regions of altered cell fate in the blade, accompanied by elimination of ligule and auricle at their wild-type positions and development of ligule and auricle in the blade at the borders of the altered regions. The affected blade cells are transformed into sheath-like cells, as determined by morphological and genetic tests. Lg4-O expressivity is highly dependent on genetic background. For example, two different backgrounds nay specify converse patterns of phenotypic expression. Lg4-O expressivity is also affected by the heterochronic mutation Teopod2 (Tp2). Gene dosage experiments indicates Lg4-O is a neomorph. Interactions between recessive lg mutations (which eliminate ligular structures) and the dominant Lg mutations suggest that the lg+ genes act after the Lg mutations. Lg3-O and Lg4-O act semidominantly, and interact with each other and with other mutations in the Knotted1 (Kn1)-like family (a family in which dominant mutant alleles cause blade to sheath transformation phenotypes). These interactions suggest that the above Kn1-like mutations may function similarly in the leaf. We discuss the similarities between the Lg mutations and the other mutations of the Kn1-like family, which led us to postulate that Ig3 and Ig4 are members of a growing family of kn1-like (knox) homeobox genes that are identified by dominant mutant alleles causing leaf transformation phenotypes. We also propose that certain key characteristics of this family of dominant neomorphic mutations are important for generating meaningful morphological changes during evolution.