UNIT V - CYTO SKELETON SYSTEMS

Cytoskeleton:
• The cytosol of cells contains fibers that help to maintain cell shape and mobility and
that probably provide anchoring points for the other cellular structures.
• Collectively, these fibers are termed as the cytoskeleton.
• The cytoskeleton gives cells structure and shape and allows them to move around. It’s
also important for intracellular transport.
• At least three general classes of such fibers have been identified in eukaryotic
cells. Each of these filaments is a polymer.
• All three filament systems are highly dynamic, altering their organization in response
to the needs of the cell.

A. Microtubules
•The thickest are the microtubules (20 nm in diameter) which consist primarily of the
tubulin protein.
• Each tubulin subunit is made up of one alpha and one beta-tubulin that are attached to
each other, so technically tubulin is a heterodimer, not a monomer. Since it looks like
a tube, it is named as microtubule.
• In a microtubule structure, tubulin monomers are linked both at their ends and along
their sides (laterally). This means that microtubules are quite stable along their
lengths.
• Since the tubulin subunits are always linked in the same direction, microtubules have
two distinct ends, called the plus (+) and minus (-) ends.
• On the minus end, alpha-tubulin is exposed, and on the plus end, beta-tubulin is
exposed.
• Microtubules preferentially assemble and disassemble at their plus ends.
Functions
1. Transportation of water, ions or small molecules.
2. Cytoplasmic streaming (cyclosis).
3. Formation of fibers or asters of the mitotic or meiotic spindle during cell division.
4. Formation of the structural units of the centrioles, basal granules, cilia, and flagella.

B. Microfilaments
• The thinnest are the microfilaments (7 nm in diameter) which are solid and are
principally made of two intertwined strands of a globular protein called actin. For this
reason, microfilaments are also known as actin filaments.
• Actin is powered by ATP to assemble its filamentous form, which serves as a track
for the movement of a motor protein called myosin.
 •This enables actin to engage in cellular events requiring motion such as cell division
in animal cells and cytoplasmic streaming, which is the circular movement of the cell
cytoplasm in plant cells.
Functions
1. They maintain the shape of the cell.
2. Form contractile component of cells, mainly of the muscle cells.
3. White blood cells can move to the site of an infection and engulf the pathogen due
to microfilaments.

C. Intermediate Filaments
•The fibers of the middle-order are called the intermediate filaments (IFs) having a
diameter of 10 nm.
• They are composed of a family of related proteins sharing common structural and
sequence features.
• They having been classified according to their constituent protein such as desmin
filaments, keratin filaments, neurofilaments, vimentin, and glial filaments.
Functions
1. Intermediate filaments contribute to cellular structural elements and are often
crucial in holding together tissues like skin.

D. Microtrabecular Lattice
Recently, cytoplasm has been found to be filled with a three-dimensional network of interlinked
filaments of cytoskeletal fibers, called a micro-trabecular lattice. Various cellular organelles
such as ribosomes, lysosomes, etc., are found anchored to this lattice. The micro-trabecular
lattice being flexible changes its shape and results in the change of cell shape during cell
movement.
Functions of Cytoskeleton
The cytoskeleton is responsible for lots of important cellular functions:
• In animal cells, which lack a rigid cell wall, it is the cytoskeleton that determines cell
shape.
• It allows cells to move.
• Engulf particles.
• Brace themselves against pulling forces.
• Transport vesicles through the cytosol.
• Separate chromosomes during cell division.
• It allows our muscles to contract.
Extracellular matrix

• The extracellular matrix is a mesh structure made of water and

diverse proteins and carbohydrates, which surround a cell. It serves in functions such
as cell support within a tissue, intercellular adhesion and communication, and cell
migration.
• Most cells synthesize compounds and materials destined to be secreted in the outside
space of the cell (extracellular space). These materials form an extracellular matrix
(ECM) that surrounds the cell and serves functions related to structure and
communication with the extracellular environment.
• The name extracellular matrix mainly refers to the components found outside of animal
cells. However, fungi, plant cells, and some protists have an extracellular structure
called the cell wall that some biologists consider a specialized extracellular matrix.
Other sections discuss plant cell walls in more detail; therefore, we focus on animal
cells’ extracellular matrix here.

Extracellular matrix structure

• The ECM is a network mainly composed of water and diverse proteins

and carbohydrates; the abundance of each component depends on the type of tissue they
are part of (figure 1). Some tissues are mainly formed by lots of cells with some ECM
in between (like in the brain and cardiac muscle), and others, called connective tissues,
are primarily ECM with scattered cells suspended within it.
Extracellular matrix components
As mentioned, animals’ ECM are composed mostly of water, proteins, and polysaccharides.
Variation in the way these molecules are organized and their relative amount gives a tissue its
specific texture (going from liquid and gel-like to solid), form and function. In the following,
we describe the main components found in ECMs.
A glycoprotein is a protein with one or more oligosaccharides (short chains of covalently
linked sugars) chains attached.

A proteoglycan is a molecule composed of one or more glycosaminoglycan (GAGs) chains

attached to a central protein.

A glycosaminoglycan is a long, linear polysaccharide formed by a repeating pair of sugars (for

example, hyaluronic acid, chondroitin sulfate, and heparin). They are mainly found attached to
a protein to form a proteoglycan.

Collagen
The most abundant component of animal cells’ ECM is the
glycoprotein collagen. Glycoproteins are proteins that have carbohydrates attached to them.
After exiting the cell, collagen molecules form long fibers called collagen fibrils. Collagen is
so abundant that it comprises about 30% of the proteins in animals, and as such, there are many
types of collagens. Within a tissue, collagen fibers are a mix of different types, which,
depending on the needs of the tissue, one type of collagen typically predominates over others.
The collagen fibrils organize differently depending on the tissue.
Example:
In human skin, and mammals in general, collagen fibrils form a wickerwork pattern as the skin
needs to resist pressure from multiple directions. A parallel pattern, on the other hand, like in
tendons, enables the tissue to resist tension in one major axis or direction.

Elastin
The glycoprotein elastin is also common in ECMs and associates with collagen. Elastin forms
elastic fibers that can extend, giving flexibility to tissues subjected to repeated stretch. Several
tissues that have high amounts of elastin include vasculature (circulatory system) and lung
tissues.
Example:
Elastin is present in tissues that need to be both strong and elastic (skin, blood vessels, lungs).
It is the dominant protein in arteries.

Proteoglycans
Collagen fibers are embedded in a mesh made of proteoglycans complexes. The complexes
comprise a long polysaccharide (carbohydrate) chain with hundreds
of proteoglycan molecules. The GAG chains in the proteoglycans complexes are able to
absorb high quantities of water and gives the gel-like consistency of some connective tissues.
It allows the matrix to resist compressive forces
Fibronectins and Integrins
The ECM is connected to the outside of the cell membrane by other types of receptor
glycoproteins, like fibronectins. The fibronectins bind to proteins called integrins that are
embedded into the plasma membrane. The integrins span the whole plasma membrane width,
and their cytoplasmic side binds with proteins attached to microfilaments (elements of the
cell cytoskeleton). Therefore, these glycoproteins enable cell adhesion to the ECM.
The importance of fibronectin is highlighted by the fact that mutant mice that cannot produce
this glycoprotein end up dying as embryos because endothelial cells do not form blood
vessels appropriately.

ECM component Function Examples of tissues where it is

found
Proteoglycans They fill most of the Found in all tissues, the amount
extracellular space and of water they absorb depends on
gives the hydrogel the function (for example,
consistency of tissues, resistance to compressive forces
which allows the matrix is important in cartilage).
to resist compressive
forces. It may help to
regulate the traffic of
cells and molecules
through the ECM.
Collagen (glycoprotein) Gives mechanical Found in all tissues, and it is the
support to tissues and main components in bone and
resistance to skin.
tensile/stretching forces.
The most abundant
component of animals’
ECM.
Elastin (glycoprotein) Gives flexibility to Found in many tissues like skin,
tissues. Also abundant in vasculature (circulatory system),
ECMs, usually and lung tissues.
associated with collagen.
Fibronectin (glycoprotein) Attachment to cells (they Found in all tissues.
attach to integrins in the
plasma membrane), cell
migration.
Integrin (glycoprotein) Cell attachment to the Found in all tissues.
ECM, cell
communication, signal
transmission.

Extracellular matrix function

The ECM provides mechanical support to the cells in a tissue and regulates intercellular
adhesion and communication. Recent research shows that the ECM is highly dynamic and of
vital importance. It determines and controls essential behaviors and characteristics of cells such
as proliferation, adhesion, migration, polarity, differentiation, and apoptosis.
Mechanical and structural support
The strong collagen fibers of the ECM mainly give mechanical support throughout tissues.
Additionally, the carbohydrate chains that form a proteoglycan molecule are very good for
absorbing water, and the amount of water differs for particular tissues and can give the matrix
a hydrated gel consistency. Therefore, the ECM also serves for resisting compressive forces
due to its gel consistency.
When you walk around, or run and jump, your joints have to withstand significant compressive
forces. They can do that because of cartilage, a type of connective tissue whose ECM has a
large amount of water. Thus, the resistance of ECM to compressive forces (shock absorbance)
is significant in some tissues like cartilage.

Because of its structure, the ECM can work as a physical barrier, an anchorage site, or a
movement track for cell migration. The glycoprotein fibronectin functions in cell attachment
and migration. Cell migration (movement of cells inside the body or changes in their position)
is essential in processes like wound healing and the growth of a fetus.
Regulation of cell behavior and communication
Integrins are called that way because they integrate or connect the outside and inside of the
cell. Their extracellular side binds with the fibronectins, and their cytoplasmic side with
the cytoskeleton; they serve functions in cellular communication and signal
transmission. Matrix components can activate integrins and change their conformation, which
is important, because protein function is directly related to their conformation.

The activated integrin then initiates signaling pathways that affect cell proliferation,
differentiation, polarity, contractility, and gene expression. On the other hand, intracellular
signals can also activate integrins and trigger signaling to the outside.

Cells can sense the biochemical properties of the ECM, including growth factors and
bioactive molecules, and interact accordingly with the outside environment. However, cells
can also sense the physical properties of the matrix, like rigidity, density, porosity, and
insolubility.
For example, stem cells surrounded by a softer matrix usually follow neurogenesis, and when
surrounded by a stiffer matrix, they follow osteogenesis, which is the process in which bone
cells are generated.