Coordination of spindle orientation with the axis of cell division is an es
sential process in all eukaryotes. In addition to ensuring accurate chromos
omal segregation, proper spindle orientation also establishes differential
cell fates and proper morphogenesis(1). In both animal and yeast cells, thi
s process is dependent on cytoplasmic microtubules interacting with the cor
tical actin-based cytoskeleton(2-5), although the motive force was unknown.
Here we show that yeast Myo2, a myosin V that translocates along polarized
actin cables into the bud(6), orientates the spindle early in the cell cyc
le by binding and polarizing the microtubule-associated protein Kar9 (refs
7-9). The tail domain of Myo2 that binds Kar9 also interacts with secretory
vesicles(12) and vacuolar elements(13), making it a pivotal component of y
east cell polarization.