  The growth of actin filament networks is a fundamental biological process
that drives a variety of cellular and intracellular motions. During motility,
eukaryotic cells and intracellular pathogens are propelled by actin networks
organized by nucleation-promoting factors, which trigger the formation of
nascent filaments off the side of existing filaments in the network. A Brownian
ratchet (BR) mechanism has been proposed to couple actin polymerization to
cellular movements, whereby thermal motions are rectified by the addition of
actin monomers at the end of growing filaments. Here, by following
actin--propelled microspheres using three--dimensional laser tracking, we find
that beads adhered to the growing network move via an object--fluctuating BR.
Velocity varies with the amplitude of thermal fluctuation and inversely with
viscosity as predicted for a BR. In addition, motion is saltatory with a broad
distribution of step sizes that is correlated in time. These data point to a
model in which thermal fluctuations of the microsphere or entire actin network,
and not individual filaments, govern motility. This conclusion is supported by
Monte Carlo simulations of an adhesion--based BR and suggests an important role
for membrane tension in the control of actin--based cellular protrusions.
