Molecular regulation of starch metabolism

Abstract

Starch is the second most abundant biomass next to cellulose and composed of amylopectin, a highly branched glucan, and amylose, an essentially linear glucan. The former and the latter glucans usually account for approximately 65–85% and 15–35% of the total starch, respectively. During the last three decades the basic scheme of starch biosynthesis has been established based on numerous biochemical, genetic, and molecular biological approaches worldwide using a variety of higher plants and algae. It is well known that after the synthesis of ADPglucose by ADPglucose pyrophosphorylase (AGPase), amylopectin’s fne structure is formed by concerted actions of multiple isozymes from three classes of enzymes, starch synthase (SS), starch branching enzyme (BE), and starch debranching enzyme (DBE), and that amylose is synthesized by mainly granule-bound SS (GBSS). In addition to the roles of starch biosynthetic isozymes, the contributions of α-glucan phosphorylase, α-glucan, water dikinase, phosphoglucan, water dikinase, pyruvate, phosphate dikinase, α-amylase, and carbohydrate-binding modules have been documented. Information on the whole genome sequence and omics analyses are available in main plant species. All these results revealed the roles of key biosynthetic isozymes of SS, GBSS, BE, and DBE and subunits of AGPase to starch biosynthesis, and presently we know to what extent the fne structure of starch molecules and the internal structure and physicochemical properties of starch granules as well as starch amounts can be modifed in accord with the activity levels of these isozymes and subunits. However, in spite of numerous past investigations, the regulation of the network of enzymatic reactions has not been fully understood. To resolve the complex mechanisms, we need to examine several topics such as redundancy and supplementary functions of multiple isozymes, enzymeenzyme interaction(s), and regulatory factors controlling catalytic and specific activities of individual isozymes, temporal and spatial co-expression of multiple isozymes, post-translational modifcation of enzymatic capacities such as phosphorylation, glycosylation, and redox state. There are still lots of uncertainties in the understanding of the initiation of starch biosynthesis.