We measured the lengths of actin filaments formed by spontaneous polymeriza
tion of highly purified actin monomers by fluorescence microscopy after lab
eling with rhodamine-phalloidin. The length distributions are exponential w
ith a mean of similar to 7 mu m (2600 subunits). This length is independent
of the initial concentration of actin monomer, an observation inconsistent
with a simple nucleation-elongation mechanism. However, with the addition
of physically reasonable rates of filament annealing and fragmenting, a nuc
leation-elongation mechanism can reproduce the observed average length of f
ilaments in two types of experiments: 1) filaments formed from a wide range
of highly purified actin monomer concentrations, and 2) filaments formed f
rom 24 mu M actin over a range of CapZ concentrations.