The entry of carbon from sucrose into cellular metabolism in plants can potentially be catalyzed by either sucrose synthase (SUS) or invertase (INV). These 2 routes have different implications for cellular metabolism in general and for the production of key metabolites, including the cell-wall precursor UDPglucose. To examine the importance of these 2 routes of sucrose catabolism in Arabidopsis thaliana (L.), we generated mutant plants that lack 4 of the 6 isoforms of SUS. These mutants (sus1/sus2/sus3/sus4 mutants) lack SUS activity in all cell types except the phloem. Surprisingly, the mutant plants are normal with respect to starch and sugar content, seed weight and lipid content, cellulose content, and cell-wall structure. Plants lacking the remaining 2 isoforms of SUS (sus5/sus6 mutants), which are expressed specifically in the phloem, have reduced amounts of callose in the sieve plates of the sieve elements. To discover whether sucrose catabolism in Arabidopsis requires INVs rather than SUSs, we further generated plants deficient in 2 closely related isoforms of neutral INV predicted to be the main cytosolic forms in the root (cinv1/cinv2 mutants). The mutant plants have severely reduced growth rates. We discuss the implications of these findings for our understanding of carbon supply to the nonphotosynthetic cells of plants.
Most plant cells receive essentially all of their carbon as sucrose. Sucrose catabolism in plants is one of the largest metabolic fluxes on the planet, second only to fluxes in primary carbon assimilation. Only 2 enzymes can catalyze sucrose catabolism under physiological conditions: sucrose synthase (SUS) and invertase (INV); thus, most plant biomass is derived via 1 of these 2 routes. However, despite their central role in carbon partitioning and biomass accumulation, the precise roles and relative importance of these enzymes remain largely unknown.SUS and INV both occur as multiple, distinct isoforms. INV catalyzes the effectively irreversible hydrolysis of sucrose to glucose and fructose. Isoforms in the cell wall and vacuole (acid INV) differ in structure from those predicted to be in the cytosol, mitochondria and plastids (neutral/alkaline INV). SUS catalyzes the reversible conversion of sucrose to fructose and UDPglucose; SUS isoforms are believed to be cytosolic.Several lines of evidence indicate a predominant role for SUS in the entry of carbon into metabolism in nonphotosynthetic cells. Individual isoforms are needed for normal development in some plant organs, including potato tuber, pea and maize seed, tomato fruit, and cotton fibers (1-5). SUS is held to be important in determining sink strength, and in phloem loading (1, 6, 7). It is also proposed to have specific roles in cellulose synthesis, and in starch synthesis in leaves. In the widely cited model for cellulose synthesis, the substrate UDPglucose is channeled to the cellulose synthase complex in the plasma membrane via a SUS associated with the inner face of the complex (8, 9). Consistent with this idea, some SUS a...