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Biophys J, January 2001, p. 149-154, Vol. 80, No. 1
*Department of Chemistry, Rice University, Houston, Texas
77005-1892; and
Institute for Physical Science and
Technology, University of Maryland, College Park, Maryland 20742 USA
Forces generated by microtubule polymerization and
depolymerization are important for the biological functioning of cells. The mean growth velocity, V, under an opposing force,
F, has been measured by Dogterom and Yurke (1997;
Science 278:856-860) for single microtubules growing in
vitro, but their analysis of the data suggested that V
decreased more rapidly with F than equilibrium (or
"thermodynamic") theory predicted and entailed negative values for
the dissociation rate and undefined (or unreasonable) values for the
stall force, at which V vanishes. By contrast,
considering the mean work done against the external load and allowing
for load-distribution factors for the "on" and "off" rates, we
find good agreement with a simple theory that yields a plausible
stalling force. Although specific numerical results are sensitive to
choice of fitting criteria, about 80% of the variation with load is
carried by the "off" (or dissociation) rate, but, since that is
small (in accordance with independent observations), the dominant force dependence comes from the "on" rate, which is associated with a
displacement length, d1, significantly
longer than d0
1/13(8.2 nm), the
mean length increase per added tubulin dimer. Measuring the dispersion
in length of the growing microtubules could provide a check. The theory
implies that the stationary stall state (at V = 0)
is not one of simple associative thermal equilibrium, as previously
supposed; rather, it appears to be dissipative and kinetically controlled.
Biophys J, January 2001, p. 149-154, Vol. 80, No. 1
© 2001 by the Biophysical Society 0006-3495/01/01/149/06 $2.00
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