p. 322 free
energy source an enzyme can only catalyze a reaction towards but not against
the equilibrium. If the product dissociation step is thought irreversible,
then whether the S to P conversion is energetically favorable or not, the
reaction will proceed clockwise anyway. We know from our experience with
hundreds of enzymes that this is not the case. The inconsistency can easily
be be removed by writing a reversible product dissociation step. Now the
reaction becomes scheme 5. Scheme 5 has been used to describe many membrane
transport and energy transducing processes. The reaction of scheme 5 can
proceed clockwise or counter-clockwise depending on the sign of the free
energy of the S to P conversion, clockwise if (delta)G is negative and
counter-clockwise if it is positive. More importantly, when a cell membrane
is involved this scheme represents an efficient mechanism for energy and
signal transductions as we shall see later; yet, the enzyme E is basically
a Michaelis-Menten enzyme. An external energy source can be coupled to
this scheme so that enzyme can catalyze a reaction against its chemical
potential gradient. Scheme 5 has a characteristic frequency. Only an oscillating
driving force the frequency of which matches this charateristic frequency
will be effective in propelling the "catalytic wheel" of scheme 5, as will
be discussed more explicitly in the following sections.