Balancing division and expansion during maize leaf morphogenesis: analysis of the mutant, warty-1.

Cell division and expansion are growth events that contribute to the developing shape, or morphogenesis, of a plant. Division and expansion are coordinated to the extent that plant organs, such as leaves, generally portray a predictable cellular pattern. To dissect the relationship between division and expansion, and to test for the role of each during morphogenesis, we have identified a recessive mutation warty-1 that produces a primary defect in cell size and shape in mutant leaves. Warty-1 mutant plants are similar to non-mutant siblings in terms of flowering time, overall plant size and leaf shape. Mature adult leaves have raised warts, consisting of excessively enlarged cells, that appear in patchy distribution throughout the blade. Cell wall deposition is abnormal or incomplete, suggesting cytokinesis is also affected, either directly or indirectly. Cells first increase in size at specific positions, which correspond to predictable cell dimensions of a developing 1 cm leaf. Once mutant cells exceed 133% normal size, cytokinesis becomes abnormal. As differentiation progresses, cells that appear normal in the mutant are actually dividing faster and are smaller than comparable cells in non-mutant siblings. These results suggest that (1) cells may compensate for growth defects by altering their cell cycle and that (2) proper execution of cytokinesis may require that cell size ratios are properly maintained.