Cytoskeleton
Eucaryotic cells contain protein fibers that are involved in
- establishing cell shape
- providing mechanical strength
 - cell movement
 - chromosome separation
-intracellular transport of organelles
• There are three main types of fibers in the
  cytoskeleton:
• microtubules (25 nm diameter),
• microfilaments (actin filament: 5-9 nm),
•
• intermediate filaments (10 nm).
                      Microtubules
An a,b-tubulin heterodimer is the basic structural unit of microtubules. The
heterodimer does not come apart, once formed. The a and b tubulins, which
are each about 55 kDa MW, are homologous but not identical. Each has a
nucleotide binding site. a-Tubulin has a bound molecule of GTP, that does not
hydrolyze. b-Tubulin may have bound GTP or GDP.
A microtubule is a hollow cylinder about 25 nm in diameter. Along the
microtubule axis, tubulin heterodimers are joined end-to-end to
form protofilaments, with alternating a & b subunits. Staggered assembly
of 13 protofilaments yields a helical arrangement of tubulin heterodimers in
the cylinder wall.
GTP must be bound to both a and b subunits for a tubulin heterodimer to
associate with other heterodimers to form a protofilament or microtubule.
Subunit addition brings b-tubulin that was exposed at the plus end into contact
with a-tubulin. This promotes hydrolysis of GTP bound to the now
interior b-tubulin. Pi dissociates, butb-tubulin within a microtubule cannot
exchange its bound GDP for GTP. The GTP on a-tubulin does not hydrolyze.
Microtubules
               The structure of a microtubule and its subunit
The tubulin subunits assemble head-to-tail to create polar
filaments
Microtubule Motor Protein
• Cilia usually occur in large numbers on the
  cell surface.
  – They are about 0.25 microns in diameter and
    2-20 microns long.
• There are usually just one or a few flagella
  per cell.
  – Flagella are the same width as cilia, but 10-200
    microns long.
• In spite of their differences, both cilia and
  flagella have the same ultrastructure.
  – Both have a core of microtubules sheathed by
    the plasma membrane.
  – Nine doublets of microtubules arranged around
    a pair at the center, the “9 + 2” pattern.
  – Flexible “wheels” of proteins connect outer
    doublets to each other and to the core.
  – The outer doublets are also connected by
    motor proteins.
  – The cilium or flagellum is anchored in the cell
    by a basal body, whose structure is identical to
    a centriole.
• The bending of cilia and flagella is driven by
  the arms of a motor protein, dynein.
  – Addition to dynein of a phosphate group from
    ATP and its removal causes conformation
    changes in the protein.
  – Dynein arms alternately
    grab, move, and release
    the outer microtubules.
  – Protein cross-links limit
    sliding and the force is
    expressed as bending.
                            Microfilament
Microfilaments – actin filaments.
They are built from molecules of a globular protein – actin.
                  Role of Microfilament
Changes in cell shape
Muscle contraction
Cytoplasmic streaming
Cell motility
Cell division – cleavage furrow
formation
• In muscle cells, thousands of actin filaments are arranged parallel to
  one another.
• Thicker filaments, composed of a motor protein, myosin, interdigitate
  with the thinner actin fibers.
   – Myosin molecules walk along the actin filament, pulling stacks of
      actin fibers together and shortening the cell.
• In plant cells (and others), actin-myosin
  interactions and sol-gel transformations
  drive cytoplasmic streaming.
  – This creates a circular flow of cytoplasm in the
    cell.
  – This speeds the distribution of materials within
    the cell.
       Fig. 7.21c