The expression of a sucrose-phosphate synthase (SPS) gene from maize (Zea mays, a monocotyledon) in tomato (Lycopersicon esculenfum, a dicotyledon) resulted i n marked increases in extractable SPS activity in the light and the dark. Diurna1 modulation of the native tomato SPS activity was found. However, when the maize enzyme was present the tomato leaf cells were unable to regulate its activation state. No detrimental effects were observed and total dry matter production was unchanged. However, carbon allocation within the plants was modified such that in shoots it increased, whereas in roots it decreased. There was, therefore, a change in the shootroot dry weight ratio favoring the shoot. This was positively correlated with increased SPS activity in leaves. SPS was a major determinant of the amount of starch i n leaves as well as sucrose. There was a strong positive correlation between the ratio of sucrose to starch and SPS activity in leaves. Therefore, SPS activity is a major determinant of the partitioning of photosynthetically fixed carbon in the leaf and in the whole plant. The photosynthetic rate in air was not significantly increased as a result of elevated leaf SPS activity. However, the light-and C02-saturated rate of photosynthesis was increased by about 20% in leaves expressing high SPS. In addition, the temporary enhancement of the photosynthetic rate following brief exposures to low light was increased in the high SPS plants relative to controls. We conclude that the leve1 of SPS in the leaves plays a pivotal role in carbon partitioning. Furthermore, high SPS levels have the potential to boost photosynthetic rates under favorable conditions. ~ ~~~~~~~~ SPS catalyzes the penultimate step of sucrose biosynthesis in leaves (Pontis, 1977). This rate-limiting, allosteric enzyme forms a major control point subject to regulation by metabolites and by covalent modification involving protein phosphorylation (Harbron et al., 1981; Huber, 1983;Stitt et al., 1987;Huber et al., 1989;Stitt and Quick, 1989). The activity of this enzyme can be a limiting factor for de novo SUC synthesis and also photosynthesis (Pontis, 1977; Huber, 1983;Stitt et al., 1987). The amount of extractable SPS activity is known to vary in response to light/dark transitions (Huber et al., 1987) and to the metabolic status of the leaf tissue in terms of substrate concentrations (Harbron et al., 1981; Huber, 1983; Kalt-Torres and Huber, 1987;Kerr and Huber, 1987;Stitt et al., 1987).* Corresponding author; fax 33-30-83-30-96. Multiple forms of SPS have been observed in leaves of plants such as maize (Kalt-Torres et al., 1987). The multiplicity of enzyme forms appears to arise from posttranslational modifications (Huber et al., 1989; Huber and Huber, 1991) to the enzyme protein because the cDNA isolated from maize appears to have only one predominant species of transcript (Worrell et al., 1991). The maize leaf SPS has been shown to be subject to light/dark regulation (Kalt-Torres and Huber, 1987), a process mediated by phosphorylation ...