Structure of a stem-derived callus of Ruta graveolens: meristems, leaves, and secretory structures

Abstract
Differentiation in a stem-derived callus of Ruta graveolens was studied by correlated light and electron microscopy. Shoot apical meristems, some of which initiated leaves, differentiated randomly at the surface of the callus. Some of the apical meristems had a tunica–corpus organization. Cells of the tunica and corpus had large nuclei with prominent nucleoli, small vacuoles, mitochondria, endoplasmic reticulum cisternae, and leucoplasts. The leaves were radial and had well developed stomata and chloroplasts of two types, one storing large starch grains and the other with no starch but with well developed grana. Lysigenous and schizolysigenous secretory glands were initiated in the leaves and towards the periphery of the callus. Central cells in the lysigenous glands underwent lysis forming a gland lumen into which lipid-like material from the degenerating cells was released. During early stages of cell lysis, breakdown of the middle lamella occurred, followed by the degeneration of cell walls. The lipid-like deposits are thought to be the essential oils known to be produced by these callus cultures. Schizolysigenous glands are formed by the separation of gland cells along the middle lamella and the subsequent development of an epithelial layer or layers surrounding a gland lumen. The cytology of epithelial cells was characterized by numerous ribosomes and the presence of plastids with lipid-like deposits, rough endoplasmic reticulum which occurred either as sheets of cisternae or tubular profiles, and osmiophilic deposits in the cytoplasm. Changes in the epithelial cell walls bordering on the gland lumen indicated that these cells were probably undergoing lysis in older stages of gland development.

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