COMPARATIVE BUNDLE SHEATH AND MESOPHYLL DIFFERENTIATION IN THE LEAVES OF THE C-4 GRASSES PANICUM-EFFUSUM AND PANICUM-BULBOSUM

The two C4 Panicum species examined differ in C4 acid decarboxylation type and in developmental origin of bundle sheaths in major veins of their leaf blades. In Panicum effusum R. Br. (NAD-malic enzyme type) both mesophyll (PCA) and chlorenchymatous bundle sheath (PCR) cells are derived from ground meristem. In contrast, in Panicum bulbosum H.B.K. (NADP-malic enzyme type), bundle sheath cells are derived from procambium, while mesophyll develops from ground meristem. To test the hypothesis that the developmental divergence of bundle sheath and mesophyll cells would occur earlier when these two tissues had different ontogenetic origins (in P. bulbosum) than when these tissues had the same origin (in P. effusum), the development of major veins in each species was investigated. We measured cell length and cross sectional area, plastid and mitochondrial number, plastid area, vacuole area fraction, wall thickness and fraction adjacent to intercellular space using direct and digitizer measurements of transmission electron micrographs of leaf cross sections of successive developmental stages. Many of the statistically significant changes in the structural parameters measured occurred late in development of both species. The magnitude of developmental change in P. effusum PCR cells was sometimes more dramatic, viz. changes in cell and PCR plastid area, and in mitochondrion number per cell. However, earlier divergence of PCR and PCA length and volume, and wall fraction adjacent to intercellular space in P. bulbosum than in P. effusum indicates that ontogenetic origin of PCR cells from procambium could determine the timing of at least some developmental events.