
The 26S proteasome is composed of two large subcomplexes.
The 20S core is barrel shaped, and proteolysis takes place inside
its chamber. The 19S cap are regulatory subcomplexes which can
be further divided into the base and the lid. The 19S cap is
essential for proper functioning of the proteasome; however,
its precise function is not clear. One of the subunits in the
19S cap, Rpn10 (n10), can bind polyubiquitin, suggesting that
one of the functions of the 19S cap is to bind polyubiquitinated
substrates. Several proteins in the base are ATPases (those
labeled with a t), which are thought to play a role in denaturing
protein substrate such that they can enter the narrow opening
of the catalytic core. Securin and mitotic cyclin are two of
the best documented substrates whose levels are tightly regulated
by the proteasome. Improper degradation of securin can block
chromosome segregation because securing regulates the removal
of the "glue" protein that binds the two sister chromatids
together. Inefficient proteolysis of cyclin blocks cell division
in anaphase.