Waters, Mark T.
(2004)
Plastid tubules in higher plants: an analysis of form and function.
PhD thesis, University of Nottingham.
Abstract
Besides photosynthesis, plastids are responsible for starch storage, fatty acid biosynthesis and nitrate metabolism. Our understanding of plastids can be improved with observation by microscopy, but this has been hampered by the invisibility of many plastid types. By targeting green fluorescent protein (GFP) to the plastid in transgenic plants, the visualisation of plastids has become routinely possible. Using GFP, motile, tubular protrusions can be observed to emanate from the plastid envelope into the surrounding cytoplasm. These structures, called stromules, vary considerably in frequency and length between different plastid types, but their function is poorly understood.
During tomato fruit ripening, chloroplasts in the pericarp cells differentiate into chromoplasts. As chlorophyll degrades and carotenoids accumulate, plastid and stromule morphology change dramatically. Stromules become significantly more abundant upon chromoplast differentiation, but only in one cell type where plastids are large and sparsely distributed within the cell. Ectopic chloroplast components inhibit stromule formation, whereas preventing chloroplast development leads to increased numbers of stromules. Together, these findings imply that stromule function is closely related to the differentiation status, and thus role, of the plastid in question.
In tobacco seedlings, stromules in hypocotyl epidermal cells become longer as plastids become more widely distributed within the cell, implying a plastid density-dependent regulation of stromules. Co-expression of fluorescent proteins targeted to plastids, mitochondria and peroxisomes revealed a close spatio-temporal relationship between stromules and other organelles. Stromule and plastid fusion could not be induced under conditions which promote substantial fusion of mitochondria. Data are presented suggesting that organelles may be able to pass between cells, and an experiment was designed to test this possibility in the C4 photosynthetic cells of maize.
Inhibitor studies have shown that stromule and plastid movement is dependent on the actin cytoskeleton and the ATPase activity of myosin. An Arabidopsis gene, CHUP1, is responsible for chloroplast relocation in response to light intensity and encodes a chloroplast-localised actin-binding protein. To assess whether this protein is involved in stromule movement, CHUP1 was down-regulated with RNAi. Whilst plants with reduced CHUP1 expression exhibited a chup1 mutant phenotype, no significant effect on stromules was discovered. It was thus concluded that chloroplast relocation and stromule formation are two independent processes that employ different actin-dependent mechanisms.
It is proposed that stromules act primarily to increase the plastid surface area in response to a number of developmental and environmental factors.
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