Cystolith Development and Structure in Pilea cadierei (Urticaceae)

Cystolith formation, structure and composition have been investigatedin leaves and stem internodes of Pilea cadierei (Urticaceae)using a variety of techniques at the light and electron microscopelevels. The development of lithocysts from epidermal cells hasbeen followed. These cells are cytoplasmically similar to otherepidermal cells but possess a much more active Golgi apparatusand more numerous mitochondria. The cystolith is a spindle-shapedbody composed of concentric layers of longitudinally orientatedcellulose microfibrils associated with pectins and other cellwall polysaccharides. At maturity it is heavily impregnatedwith calcium carbonate. Some cystoliths also contain siliconand are covered in a sheath of siliceous material. Cystolithformation occurs at the tip of a peg that grows in from thelithocyst wall. Evidence from ultrastructure suggests that thelithocytst cytoplasm transports carbohydrates to the cystolithvia Golgi vesicles, and organizes the deposition of cystolithcellulose microfibrils via a system of microtubules lying beneaththe plasma membrane that envelopes the growing cystolith. Thepeg is composed of heavily staining amorphous material likethat of an apoplastically sealed cell wall. It is incapableof supporting the migration of lanthanum ions into the cystolith.We conclude that cystoliths are isolated volumes of apoplastthat act as repositories for inorganic salts, principally calciumcarbonate. We propose that calcium ions move into the lithocystprotoplast from surrounding cells and are then transported acrossthe plasma membrane boundary into the cystolith. This proposalconflicts with previous suggestions that calcium enters by diffusionthrough the peg. Cystolith, lithocyst, cell wall, calcium, silicon, cytochemistry, electron probe analysis, Pilea cadierei