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The mechanism of multivalent counterion-induced bundle formation byfilamentous actin (F-actin) is studied using a coarse-grained model andmolecular dynamics simulation. Real diameter size, helically ordered chargedistribution and twist rigidity of F-actin are taken into account in our model.The attraction between parallel F-actins induced by multivalent counterions isstudied in detail and it is found that the maximum attraction occurs betweentheir closest charged domains. The model F-actins aggregate due to thelike-charge attraction and form closely packed bundles. Counterions are mostlydistributed in the narrowest gaps between neighboring F-actins inside thebundles and the channels between three adjacent F-actins correspond to lowdensity of the counterions. Density of the counterions varies periodically witha wave length comparable to the separation between consecutive G-actin monomersalong the actin polymers. Long-lived defects in the hexagonal order of F-actinsin the bundles are observed that their number increases with increasing thebundles size. Combination of electrostatic interactions and twist rigidity hasbeen found not to change the symmetry of F-actin helical conformation from thenative 13/6 symmetry. Calculation of zero-temperature energy of hexagonallyordered model F-actins with the charge of the counterions distributed ascolumns of charge domains representing counterion charge density waves hasshown that helical symmetries commensurate with the hexagonal latticecorrespond to local minima of the energy of the system. The global minimum ofenergy corresponds to 24/11 symmetry with the columns of charge domainsarranged in the narrowest gaps between the neighboring F-actins.

Chiral Structure of F-actin Bundle Formed by Multivalent Counterions?