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C4 plants have evolved >60 times from their C3 ancestors. C4 photosynthesis requires a set of closely co-ordinated anatomical and biochemical characteristics. However, it is now recognized that the evolution of C4 plants requires fewer changes than had ever been considered, because of the genetic, biochemical and anatomical pre-conditions of C3 ancestors that were recruited into C4 photosynthesis. Therefore, the pre-conditions in C3 plants are now being actively investigated to clarify the evolutionary trajectory from C3 to C4 plants and to engineer C4 traits efficiently into C3 crops. In the present mini review, the anatomical characteristics of C3 and C4 plants are briefly reviewed and the importance of the bundle sheath for the evolution of C4 photosynthesis is described. For example, while the bundle sheath of C3 rice plants accumulates large amounts of starch in the developing leaf blade and at the lamina joint of the mature leaf, the starch sheath function is also observed during leaf development in starch accumulator grasses regardless of photosynthetic type. The starch sheath function of C3 plants is therefore also implicated as a possible precondition for the evolution of C4 photosynthesis. The phylogenetic relationships between the types of storage carbohydrates and of photosynthesis need to be clarified in the future. [ABSTRACT FROM AUTHOR]

In C4 plants, bundle sheath (BS) chloroplasts are arranged in the centripetal position or in the centrifugal position, although mesophyll (M) chloroplasts are evenly distributed along cell membranes. To examine the molecular mechanism for the intracellular disposition of these chloroplasts, we observed the distribution of actin filaments in BS and M cells of the C4 plants finger millet (Eleusine coracana) and maize (Zea mays) using immunofluorescence. Fine actin filaments encircled chloroplasts in both cell types, and an actin network was observed adjacent to plasma membranes. The intracellular disposition of both chloroplasts in finger millet was disrupted by centrifugal force but recovered within 2 h in the dark. Actin filaments remained associated with chloroplasts during recovery. We also examined the effects of inhibitors on the rearrangement of chloroplasts. Inhibitors of actin polymerization, myosin-based activities and cytosolic protein synthesis blocked migration of chloroplasts. In contrast, a microtubule-depolymerizing drug had no effect. These results show that C4 plants possess a mechanism for keeping chloroplasts in the home position which is dependent on the actomyosin system and cytosolic protein synthesis but not tubulin or light. [ABSTRACT FROM AUTHOR]