The influence of chromosome flexibility on chromosome transport during anaphase A.

The role of protein flexibility in molecular motor function has previously been studied by considering a Brownian ratchet motor that is connected to its cargo by an elastic spring, with the result that the average velocity of the motor/cargo system is increased by reducing the stiffness of the linkage. Here, we extend this investigation to the case of chromosome transport during anaphase A, in which the relevant flexibility is not primarily in the motor/cargo linkage but rather in the cargo itself, i.e., in the chromosome. We model the motor mechanism as an imperfect Brownian ratchet with a built-in opposing load and the chromosome as a collection of discrete segments linked by an elastic energy function that discretizes the potential energy of an elastic rod. Thermal fluctuations are produced in the model by random forces, as in Brownian dynamics. All of the parameters that characterize the chromosome are known or can be estimated from experimental data, as can all but one of the motor parameters, which is adjusted to give the correct transport velocity of normal-length chromosomes. With the parameters so determined, we then reproduce the experimental finding of Nicklas [Nicklas, R. B. (1965) J. Cell Biol. 25, 119-135] that chromosome speed is essentially independent of chromosome length, even though our model contains no "velocity governor." We find instead that this effect is a consequence of chromosome flexibility, as it disappears when stiffer than normal chromosomes are considered.