Photosynthetic carbon assimilation is the origin of nearly a11 organic matter on earth. In most land plants, atmospheric CO, is fixed by the enzyme Rubisco via carboxylation of a five-carbon sugar phosphate. Because the first stable products of this reaction are three-carbon compounds, this type of photosynthesis is known as C, photosynthesis. In light, Rubisco also exhibits oxygenase activity (Edwards and Walker, 1983). As an oxygenase, the enzyme is involved in photorespiration through which part of the oxygenated carbon is released as CO,. Therefore, photorespiration results in carbon loss from the leaf, and when it occurs at high rates it can severely limit carbon assimilation by C, photosynthesis.The oxygenase reaction of Rubisco is favored by low CO, concentrations in the leaf because oxygen and CO, compete for binding at the active site of the enzyme. Low CO, concentrations in the leaf prevail under dry, hot conditions. Under such conditions, plants minimize water loss from the leaves by closing their stomata, which in turn restricts CO, diffusion from the atmosphere into the leaves. Plants that undergo C, photosynthesis can effectively fix carbon at low interna1 CO, concentrations. This is achieved by CO, binding via specific reactions that yield four-carbon dicarboxylic acids (Hatch, 1987). This primary carbon fixation takes place in the mesophyll compartment, in which photosynthetic cells are closer to the leaf surface. In C, plants Rubisco is confined to the bundle-sheath compartment, in which cells surround the vascular bundles. The C, acids are transported from the mesophyll to the bundle-sheath compartment, where CO, is released for refixation by Rubisco.In essence, C, biochemistry represents a biochemical CO, pump that increases the CO, concentration in the vicinity of Rubisco and thereby reduces photorespiration.In the absence of photorespiration, the quantum yield for CO, fixation is lower for C, photosynthesis than for C, photosynthesis because extra energy is required to fuel the reactions of the C, cycle (Hatch, 1987). However, when low CO,/O, ratios trigger high rates of photorespiration in C, photosynthesis, the photosynthetic performance of C, plants is superior to that of C, plants. Therefore, a low atmospheric CO, concentration may represent the evolutionary pressure for the evolution of C, photosynthesis (Hatch, 1992). It has been suggested that C, plants evolved from C, ancestors in response to a dramatic decrease in the atmospheric CO,/ O, ratio that began about 100 million years ago (Moore, 1982; Ehleringer and Monson, 1993). Among the three biochemical subtypes of C, photosynthesis, the biochemistry of the NADP-ME group, which also includes the crop plants maize and sugar cane, is probably the best characterized. The evolution of C, photosynthesis implies the compartment-specific regulation of the enzymes involved, as well as modifications in the leaf anatomy (Hatch, 1987). We have recently shown for a number of NADP-ME plants that light harvesting of PSI is also regulated in a comp...