Molecular bureaucracy: who controls the delays? Transient times in branched pathways and their control.

Analysis of metabolic control has until now been mainly confined to systems at steady state. This includes studies of the control of "transition time", which is actually a steady-state transit time that does not refer to the transient state. In this paper we examine the control of the transition state of a metabolic pathway in the approach to a stable steady state, showing that the time needed to attain it can be decreased or increased in different branches. Our analysis only applies to branched pathways, and we discuss why similar deviations cannot occur in unbranched pathways. In systems with several branches the acceleration of some branches during the transient phase, so that they reach their steady states more quickly, occurs at the expense of others, which are thus delayed. We present theorems that describe properties of the transient variables and their control.