Myosin V walks by lever action and Brownian motion

Myosin V is a molecular motor that moves cargo along actin filaments. Its two heads, each attached to a long and relatively stiff neck, move alternately forward in a "hand-over-hand" fashion. To observe under a microscope how the necks move, we attached a micrometer-sized rod to one of the necks. The leading neck swings unidirectionally forward, whereas the trailing neck, once lifted, undergoes extensive Brownian rotation in all directions before landing on a site ahead of the leading head. The neck-neck joint is essentially free, and the neck motion supports a mechanism where the active swing of the leading neck biases the random motion of the lifted head to let it eventually land on a forward site.