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microtubule
cytoskeleton
ribosome
Intermediate
filament
microfilament ER
brief introduction of cytoskeleton
The cytoskeleton is a dynamic structure that is
responsible for whole –cell movement, changes
in cell shape, contraction of muscle cells, and
provides the machinery to move organelles
from one place to another in the cytoplasm. In
addition, cytoskeleton is the master organizer
of the cell’s cytoplasm, furnishing binding
sites for the specific localization of RNAs and
proteins that were once thought to diffuse
freely through the cytoplasm.
15nm
24-26nm
Plus end +
Tread-milling model
This polarity occurs
because of the orientation
of the tubulin subunits in
the microtubule polymer.
Minus end
-
both processes always occur more rapidly
at one end, called the plus end. The
other, less active, end is the minus end.
assembly
Cytoplasmic microtubules are labile structures that
have the capacity to undergo rapid assembly and
disassembly. This characteristic is important for
many microtubule functions. Such as cell mitosis,
cell migration and establishment of cellular
polarity.
Growing microtubules have an inherent structural
polarity. This polarity occurs because of the
orientation of the tubulin subunits in the
microtubule Polymer.
disassembly
tubulin MAP1A
1.Alkaline
微 typeI MAPs microtubule
MAP1B binding domain
Microtubule-
管
associated MAP2
2.Acidic
protein (MAP
s)
typeII MAPs MAP4 protruding
domain
Microtubine polymerized
protein ( Tua )
Microtubule-associated
protein, MAP
During microtubule assembly, tubulin also
contains regions for GTP binding, for
interacting with microtubule-associated
proteins, and for several different drug-
binding sites, Such as colchicine, vinblastine
and taxol, which play an important
modulation role in assembly and
disassembly of microtubule.
The existing form of microtubule
1
2
3 A B
13 4
12 5
11 6
7
10
9 8
doublet
singlet
A B
C
triplet
The existing form of microtubule
Singlet is one of the most familiar forms in
cytoplasm. Singlet is often affected by the
temperature, colchicum and so on.
Doublet is composed of the flagellum and
cilia. A and B microtubule built up doublet,
which is composed of 23 protofilaments.
Triplet is often viewed in centrosome, which
is composed of 33 Protofilaments .
The function of microtubule
1. Microtubules are involved in intracellular
vesicle and organelle transport.
A good analogy for visualizing this event
would be to consider a railroad: the
microtubules would serve as the tracks and
the locomotory forces responsible for
transporting the vesicular cargo would be
generated by ATPases, such as kinesin and
dynein.
An example of the role of microtubules
in transporting organelles
The function of microtubule
2. Microtubules are composed of cilia and
flagella that extend from the surfaces of several
different cell types.
Flagella and cilia
Cilia are prominent in the respiratory tract and on the apical
surface of the epithelial cells that line oviduct. The major
type of flagellated cell in humans is the spermatozoon. The
beating flagellum provides the force that allows the sperm to
swim.
shaft : thin protrusion extending from
the cell surface. shaft
纤
毛
的
整 Basal body : cylindrical structure on the cilia
体 basal plasma membrane.
结
构
shaft
B A
A B Out arm
B A dynein
Inner arm
B
质膜 A A
B C1 C2
axoneme spoke
B
轴丝 A
A Spoke head
B
A B A B Nexin
Connect filament
9X2+2
B
A
When viewed in cross section , microtubules are
arranged in a distinctive nine-plus-two array.
纤
毛
本
体
基
体 (9X3+
0)
纤毛小根
B A
B A B A
B A B A
头部
头部
The sliding-microtubule mechanism
of ciliary and flagella motility
the structure of
sperm flagellum
Central pair
Outer doublet
Dynein
Bridge
Spoke
centrosome
and centriole
Protoplasm
centrosphere ( transpare 原生质
0.16-0.26um
电镜: column body
圆柱状小体— 长度 0.16-5.6um
centriole diplosome
• located in the cytoplasm attached to the outside
of the nucleus.
• Just before mitosis, the centrosome duplicates.
• The two centrosomes move apart until they are on
opposite sides of the nucleus.
• As mitosis proceeds, microtubules grow out from
each centrosome with their plus ends growing
toward the metaphase plate. These clusters of
microtubules are called spindle fibers.
The Centrosome
protoplasm
nucleus
centriole
centrosphere
C
C B C
B
A A B
C B A A
C
A B
A
B A
Centriolar A B
C B AB C
satellite C C
centriolar annulet
(9X3+0)
The structure of centrosome in electron microscope
The centrosome is composed of a centriole pair and a
surrounding cloud of amorphous substance called the
pericentriolar material (PCM). Viewed in the transverse
section, There are 9 bundles of triplet tilting just like the
wings of windmill. Between the bundles, there are many
compacting granulars.
The ultrastructure of centrosome
The function of centrosome
and centriole
As the cell enter interphase, the centrosome
is duplicated, and at the onset of mitosis,
the daughter centrosomes migrate to
opposite sides of the nucleus where they
will serve as the mitotic spindle poles.
Along with the migration, microtubule
complex is disassembled and replaced by
the mitotic apparatus.
The origin of centriole
S phase
nucleus nucleus mitosis
• Microtubules attached to opposite sides of the dyad shrink or
grow until they are of equal length.
• Microtubules motors attached to the kinetochores move them
• toward the minus end of shrinking microtubules (a dynein);
• toward the plus end of lengthening microtubules (a kinesin).
• The chromosome arms use a different kinesin to move to the
metaphase plate
Part two microfilaments
All eukaryotic cells appear to contain filaments 8
nm in diameter, called microfilaments, that are
polymers of actins.
G-actin ( globular actin monomers ) -actin
actin -actin
-actin
F-actin ( polymerization of G-actin)
-
G-actin
+
F-actin
In physiological conditions, actin monomers (G-
actin) spontaneously self associate to form
microfilaments (F-actin). This polymerised form
of actin is in constant equilibrium with G-actin.
Monomers are able to add to the ends of
filaments, form nuclei with another two
monomers to create new filaments, and to leave
filaments. The polarity of actin subunits within
microfilaments means that the two ends are
topologically different, the narrow face of actin
subunits is exposed at the pointed end while the
more bulbous face containing the NH2 and
COOH termini is exposed at the barbed end.
Pure actin can be switched between these states
in the test-tube by altering the salt concentration.
Actin in low salt conditions is in the G-state,
while adding salts causes the actin to polymerise.
These filaments , referred to as filamentous or
F-actin, are built from polymerization of a
globular actin monomer, called G-actin, which
has a relative molecular mass of 43kDa. Each
F-actin microfilament appears as two helically
intertwined chains of G-actin monomers, for
which a complete turn of the helix occurs over
a distance of 36 nm or 13 G-actin monomers.
subunits eclipse each other at what appears to
be a crossover.
microfilments
Microfilament-associated protein
Protein function
Tropomyosin stabilizes filaments
Fimbrin bundles filaments
Myosin II slides filaments in muscle
Profilin binds actin monomers
Cap Z caps plus-ends of filaments
The function of microfilament
L12 L2
L1
连 连 连
N- 端 1A
接
区 1B
接
区 2A
接
区 2B C- 端
- 螺旋 - 螺旋 - 螺旋 - 螺旋
35 8-14 101 8 9 16-17 121
The structure of intermediate filaments
All IF proteins contains a subunit-spacific
NH2-terminus of variable size, a homologous
central a-helical region of approximately
310 amino acids, and a subunit-specific
COOH-terminus of variable size. Only the
homologous 310-amino acid a-helical region is
a portion of the 10-nm IF core. The variable
regions extend from the core and are
responsible for cross-linking intermediate
filaments to other cytoskeletal structures.
Assembly of intermediate filaments
COOH 单体
NH2
mononor
COOH
NH2 二聚体
COOH
NH2 超螺旋
dimer
COOH
NH2
COOH
NH2
COOH NH2
四聚体
COOH NH2
tetramer
原丝
原丝
八聚体
原纤维
八聚体
中等纤维
中等纤维
Assembly of intermediate filaments
In the formation of the intermediate filaments,
the first step is that the 310-amino acid a-helical
region of two monomers wind around each other
into a parallel coiled-coil. The IF proteins
contain the heptad repeat within the 310-amino
acid a-helical region required for coiled-coil
formation. Next two dimers link side-by-side in
an antiparallel conformation to form a tetramer.
The IF tetramers attach laterally to each other in
a staggered array, until there are eight tetramers (
32 monomers) making up the wall of IF. The
eight tetramers are wound to form the ropelike
structure of IF.
Part three intermediate filaments
微丝 (microfilament)
中等纤维
(intermediate filament)
The camparison of cytoskeleton
microtubule microfilament Intermediate
filament
component tubulin actin Intermediate
filament monomer
1. Respiratory chain
2. Signal peptide
3. The cytoskeleton