Cytoskeleton 13.4.20

Cytoskeleton
Eucaryotic cells contain protein fibers that are involved in

Function
structural support
maintains shape of cell
provides anchorage for organelles
motility
cell locomotion
cilia, flagella, etc.
regulation
organizes structures &
activities of cell
In addition, a large number of accessory proteins, including the motor proteins,
are required for the properties associated with each of these filaments
Each type of filament has distinct mechanical properties and dynamics, but
certain fundamental principles are common to all.
Cytoskeleton Structure
network of fibers extending throughout cytoplasm
3 main protein fibers
Microtubules
Microfilaments
intermediate filaments
Evolutionary perspective
Proteins that make up the fibers are very similar in all

living things
from bacteria to humans
tubulin (all cells)

actin (eukaryote cells)
Means that they are both ancient and essential for life
There are three main types of fibers in the
cytoskeleton:
1. Microfilaments (actin filament: 5-9 nm),

2. Microtubules (25 nm diameter),
3. Intermediate filaments (10 nm).
Microfilaments
Some functions of actin filaments are:

- to provide mechanical strength to the cell by forming a band under the plasma membrane
- link transmembrane proteins to cytoplasmic proteins
- form contractile ring during cytokinesis in animal cells
- cytoplasmic streaming
- generate locomotion in cells such as white blood cells and amoeba
- Interact with myosin to provide force of muscular contraction
Microtubules participate in a wide variety of cell activities. Most involve motion that
is provided by protein “motors” that use ATP. They determine the positions of
membrane-enclosed organelles and direct intracellular transport. The migration of
chromosomes during mitosis and meiosis takes place on microtubules that make up
the spindle fibers.
Intermediate filaments provide mechanical strength and resistance to shear stress.
There are several types of intermediate filaments, each constructed from one or
more proteins characteristic of it.
Keratins are found in epithelial cells, hair and nails

Nuclear lamins form a meshwork that stabilizes the inner nuclear membrane
Neurofilaments strengthen the long axons of neurons
Vimentins provide mechanical strength to muscle and other cells
Microfilaments
Molecular structure of actin
G-actin F-actin
( Garland Science Molecular Biology of the Cell 2008)

Microfilament
Microfilaments – actin filaments.

They are built from molecules of a globular protein – actin.
Actin and accessory proteins
Molecular Biology of the Cell (© Garland Science 2008)

Actin binding proteins (ABPs)
3 groups:
 banding and cross linking proteins

 regulatory proteins: Organisation of
polymerization/depolymerization, actin filaments
severing proteins,capping proteins
 Motor proteins
- sliding on MF (myosin) Sliding
Cross-linking proteins I.
Contractile bundle Parallel bundle

Actin and muscle movements
• 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.
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.
Microtubules
The structure of a microtubule and its subunit
The tubulin subunits assemble head-to-tail to create polar filaments

Figure 1. Conventional kinesin motor proteins translocate along the microtubules in the positive
direction, while the dynein motors step along the microtubules in the opposite direction.
kinesin
- +
dynein
cAMP cAMP
pigment cells
Microtubular systems in the cells
-Centrosome Interphase cell
centrosome
-Cilia / flagellum
Cilla
Basal body
Dividing cell spindle
-Mitotic system
Neuron
centrosome
- Vesicular transport axon

• Microtubules of mitotic spindle and kinetochore
How motor proteins can organise the position of cell
organelles (ER, Golgi) ?
(Hirokawa, N. Science 1998, 279:519

Cilia and Flagella
Role of the dynein arms in beating cilia
Telescopic effect Beating

• Intermediate filaments
Structure of an intermediate filaments
Monomer
Parallel dimer
Antiparallel
tetramer
Protofilaments
Intermedaite
filaments

Mechanical characterization of cytoskeleton components
intermedier filament
i.e. vimentin
microtubule
deformation
= rupture
actin filament
force
Role of intermedier filaments
Buffer against external mechanical stress
Tissue specificity !!!
Epithel – keratin
Vimentin
Connective tissue
Muscles
Neuroglia
} vimentin-like
Desmin
Glial protein
Neurones(axon) - neurofilaments
Exception:
Nucleus – lamines (A,B,C) →(lamina fibrosa)

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Cytoskeleton

Eucaryotic cells contain protein fibers that are involved in

network of fibers extending throughout cytoplasm

3 main protein fibers

Proteins that make up the fibers are very similar in all

from bacteria to humans

tubulin (all cells)

1. Microfilaments (actin filament: 5-9 nm),

Some functions of actin filaments are:

Keratins are found in epithelial cells, hair and nails

( Garland Science Molecular Biology of the Cell 2008)

Microfilaments – actin filaments.

Molecular Biology of the Cell (© Garland Science 2008)

 banding and cross linking proteins

Contractile bundle Parallel bundle

Molecular Biology of the Cell (© Garland Science 2008)

A microtubule is a hollow cylinder about 25 nm in diameter. Along the

GTP must be bound to both a and b subunits for a tubulin heterodimer to

The tubulin subunits assemble head-to-tail to create polar filaments

-Centrosome Interphase cell

Dividing cell spindle

- Vesicular transport axon

(Hirokawa, N. Science 1998, 279:519

Telescopic effect Beating

Molecular Biology of the Cell (© Garland Science 2008)

Buffer against external mechanical stress

Tissue specificity !!!

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