Relationships between enzymatic flux capacities and metabolic flux rates: Nonequilibrium reactions in muscle glycolysis

The rules that govern the relationships between enzymatic flux capacities (V-max) and maximum physiological flux rates (nu) at enzyme-catalyzed steps in pathways are poorly understood, We relate in vitro V-max values with in vive flux rates for glycogen phosphorylase, hexokinase, and phosphofructokinase, enzymes catalyzing nonequilibrium reactions, from a variety of muscle types in fishes, insects, birds, and mammals, Flux capacities are in large excess over physiological flux rates in low-flux muscles, resulting in low fractional velocities (%V-max = nu/V-max x 100) in vivo, In high-flux muscles, close matches between flux capacities and flux rates (resulting in fractional velocities approaching 100% in vivo) are observed, These empirical observations are reconciled with current concepts concerning enzyme function and regulation, We suggest that in high-flux muscles, close matches between enzymatic flux capacities and metabolic flux rates (i.e., the lack of excess capacities) may result from space constraints in the sarcoplasm.