In biomolecular systems, the mechanical transfer of free energy occurs with
both high efficiency and high speed, It is shown here that such a transfer
can be achieved only if the participating free-energy-storing elements exh
ibit opposing relationships between their content of free energy and the fo
rce they exert in the transfer direction, A kinetic equilibrium of forces (
KEF) results, in which the transfer of free energy is mediated essentially
by thermal molecular motion, On the basis of present evidence, KEF is used
as a guiding principle in developing a mechanical model of the crossbridge
cycle in muscle contraction. The model allows the basic features of molecul
ar events to be visualized in terms of plausible structures, Real understan
ding of the process will require identification of the elements that perfor
m the functions described here. Besides chemomechanical energy transduction
, KEF may have a role in other biomolecular processes in which free energy
is transferred mechanically over large distances.