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Torque force in the kinesin stroke drives coupled yaw axis and orbital rotations of kinesin coated gold nanorods

Kinesin motor domains generate impulses of force and movement that have both translational and rotational components, raising the question of how the rotational component contributes to motor function. We used a new assay in which kinesin-coated gold nanorods (kinesin-GNRs) move on suspended microtubules, for three plus-end-directed kinesins: single-headed KIF1A, dimeric ZEN-4 and single-headed kinesin-1. Polarization of the light scattered by all three types of kinesin-GNRs periodically oscillated as they orbited the microtubule along a left-handed helical trajectory. Our analyses revealed that each kinesin-GNR unidirectionally rotates about its yaw axis as it translocates, and that the period of this yaw-axis rotation corresponds to two periods of its left-handed helical orbit around the microtubule axis. Stochastic simulations suggest that the yaw-axis rotation enhances biased lateral displacement of the kinesin team. Our study reveals biaxial rotation as a new mode of motility in kinesin teams that helps the team to sidestep obstacles.