Network organization of cell metabolism: monosaccharide interconversion.

The structural properties of carbohydrate metabolism are being studied. The present contribution focuses mainly on those processes involving the transfer of carbon fragments among sugars. It is shown how enzymatic activities fix the way the system self-organizes stoichiometrically at the steady state. It is proven that there exists a specific correspondence between the set of all possible enzymic activities, the activity set, and the set of stoichiometrically compatible flux distributions through the pathway. On the one hand, there are enzymic activities that do not allow a stoichiometrically feasible coupling at the steady state of the reactions involved in the conversion. On the other hand, there are enzymic activities that are related to one or more flux distributions at the steady state (i.e. with one or several rate vectors respectively). For this latter group, it can be demonstrated that the structure of the system depends on other non-structural factors, such as boundary constraints and the kinetic parameters. As a consequence, it is suggested that this kind of metabolic process must be viewed as a complex reaction network instead of a sequential number of steps. Some implications of these derivations are illustrated for the particular conversion of CO2 --> C3. General remarks are also discussed within the framework of network models of cell metabolism.