A model for Bud6's role in directing the program
of cytoplasmic microtubule interactions during spindle assembly. (A)
After bud emergence, positional information at the bud tip directs
microtubule interactions, thereby orienting duplicated SPBs facing the
bud neck (a). As SPBs separate, the SPBdaughter retains
contacts to the bud tip (b). In concert with spindle assembly, de novo
cytoplasmic microtubules are directed to a new Bud6-dependent area of
capture at the neck and prevented from undergoing capture in the bud
(c). As spindle assembly proceeds, cytoplasmic microtubule interactions
draw the SPBmother away from the neck and cause spindle
alignment (d and f). Neck interactions also contribute to spindle
retention at the neck. Premature organization of cytoplasmic
microtubules in cdc28-4 clb5 cells (Segal et
al., 2000) favors de novo contacts with the bud tip before an
area for cytoplasmic microtubule–bud neck interactions develops. Bud6
partition between the bud tip and neck is influenced by Bni1 and Bud3,
respectively. (B) Alternative modes of spindle orientation in cortical
cue mutants. A bud6 mutation impairs initial orientation
of duplicated SPBs facing the bud neck and ensuing neck interactions.
Orientation relies primarily on microtubule search and capture by the
bud. A bni1 mutation directs interactions to the neck
throughout spindle morphogenesis. In both mutants, Kar9 still
contributes to orientation. On the other hand, a kar9
mutation reduces cytoplasmic microtubule capture in the bud while
orientation via neck interactions can occur after spindle assembly
(when Bud6 accumulates at the neck) or during anaphase.