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Biomechanical analysis of the Rolled (RLD) leaf phenotype of maize¹

¹ Section of Plant Biology and ⁴ Department of Land, Air and Water Resources, University of California, Davis, California 95616-8627 USA; ² S.E.P.F.-I.N.R.A. 86 600 Lusignan France; and ³ Department of Plant and Microbial Biology, University of California, Berkeley, California 94720-3102 USA

ABSTRACT

The pleiotropic effects of the Rld1-O/+ mutation of Zea mays (Poaceae) on leaf phenotype include a suppression of normal transverse unrolling, a reversed top/bottom epidermal polarity, and an apparently straighter longitudinal shape. According to engineering shell theory, there might be mechanical coupling between transverse and longitudinal habit, i.e., the leaf rolling itself might produce the longitudinal straightening. We tested this possibility with quantitative curvature measurements and mechanical uncoupling experiments. The contributions of elastic bending under self weight, mechanical coupling, and rest state of leaf parts to the longitudinal and transverse habit were assessed in Rld1-O/+ mutants and a population of sibling +/+ segregants. Elastic bending and curvature coupling are shown to be relatively unimportant. The Rld1-O/+ mutation is shown to alter not only the unrolling process, but also the developmental longitudinal curving in the growing leaf, leading to a straighter midrib and a rolled lamina. The Rld1-O/+ mutant is thus a suitable model to study the relation between tissue polarity and differential curvature development in the maize leaf. Since on the abaxial side of the leaf, more abundant sclerenchyma is found in +/+ than in Rld1-O/+, a gradient in sclerification may contribute to the development of midrib curvature.

Key Words: biomechanics • curvature • development • grass • leaf rolling • leaf shape • Poaceae • Rld mutant • sclerenchyma • Zea mays

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