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Cell Junctions
Specialized cell junctions occur at points of cell-cell and cell-matrix contact in all tissues, and they are
particularly plentiful in epithelia.
Cell junctions can be classified into three functional groups:
+ Occluding junctions seal cells together in an epithelium in a way that prevents even small
molecules from leaking from one side of the sheet to the other.
‘+ Anchoring junctions mechanically attach cells (and their cytoskeletons) to their neighbors or to
the extracellular matrix,
+ Communicating junctions mediate the passage of chemical or electrical signals from one
interacting cell to its partner.
+ QccLUDING JUNCTIONS
‘tight junctions (vertebrates only)
2.septate junctions (invertebrates mainly)
+ ANCHORING JUNCTIONS
+ Actin filament attachment sites
41. celkcell junctions (adherens junctions)
2. cell matrix Junctions (focal adhesions)
+ Intermediate flament attachment sites
1. celkcell junctions (desmosomes)
2. celmatrix junctions {hemidesmosomes)
+ COMMUNICATING JUNCTIONS
1. gap junctions
2. chemical synapses
3. plasmodesmata (plants only)
+ Occluding Junctions —
Tight Junction (zonuta occludens) —
+ Form a Selective Permeability Barrier Across Epithelial Cell Sheets+The tight junctions between epithelial cells have two roles.
1. Function as barriers to the diffusion of some membrane proteins (and lipids) between apical and
basolateral domains of the plasina membrane , Mixing of such proteins and lipids occurs if tight junctions
are disrupted.
2. Seal neighboring cells together so that, if @ low-molecutar-weight tracer is added to one side of an
epithetlum, tt will generally not pass beyond the tight junction.
+ Permeability to small molecules varies greatly In different epithelia.
‘+ Tight junctions in the epithelium fining the small intestine, for example, are 10,000 times more
permeable to inorganic fons, such as Na, than the tight junctions in the epithelium lining the
urinary bladder.
PROTEINS IN TIGHT JUNCTION
‘The major transmembrane proteins in a tight junction are the
+ dlaudins, which are essential for tight Junction formation and function and differ in different,
tight junetions,
A second major transmembrane protein in tight junetions Is
“ ceeludin, the function of which Is uncertain,
“ Claudins and occludins associate with Intracellular peripheral membrane proteins celled ZO
proteins (a tight junction is also known as azonuta occludens), which anchor the strands to
the actin cytoskeleton
+ Ininvertebrates, septate junctions are the main occluding junction.
The role of tight junctions in transcellular transport
In the example shown, glucose is actively transported into the cell by Na-driven glucose symports at
the apical surface, and it diffuses out of the cell by facilitated diffusion mediated by glucose carriers in
the basolateral membrane.
Tight junctions are thought to
4 confine the transport proteins to their appropriate membrane domains by acting as diffusion
barriers within the lipid bilayer of the plasma membrane;
+ these junctions also block the backflow of glucose from the basal side of the epithelium into the
gutlumenNa*-driven
glucose symport
of adjacent cells
intercellular
space
passive glucose
carrier protein
basolateral
surface,
Va
extracellular
fluid ——
BLOOD
Adherens Junction
+ Adherens junctions are also referred to as zonula adherens, intermediate junction, or as belt
desmosomes. Zonula means small zone or belt-like, and adherens refers to adhesion (sticking
together).+ Adherens junctions are built primarily from cadherins, whose extracellular segments bind to
each other and whose intracellular segments bind to catenins. Catenins are connected to actin
filaments.
+ Provides a strong bond between the sides of adjacent epithelial cell membranes,
Structure and Composition
‘The zonula adherens is composed af several different proteins:
+The actin microfilaments of the cytoskeleton (the internal skeleton of the cell}.
+ Anchor proteins, found inside each cell. These are called catenin, vineulin, and alpha-actinin,
‘They link the actin microfilaments to the cadherins.
+ Cadherins, namely E-cadherin. These are transmembrane adhesion proteins, whose main
portions are located in the extracellular space.
+ The extracellular part of one cell's cadherin binds to the extracellular part of the adjacent cell's
cadherin in the space between the two cells,
+ This intracellular (within the cell) tail then links up to catenin proteins to form the
cadherin-catenin complex. This complex binds to vinculin and alpha-actinin - these two proteins
link the cadherin-catenin complex to the cell's internal skeletal framework (the actin
microfilaments).
RGRGrenS TUNCHON] tyemenetene
Desmosomes
+ Desmosomes are a type of anchoring junction in animal tissues that connect adjacent cells.
+ Desmesomes have intermediate filaments in the cells underneath that help anchor the junction+ Pemphigus is an autoimmune disease in which the patient has developed antibodies against
proteins (cadherins) in desmosomes. The loosening of the adhesion between adjacent epithelial
cells causes blistering
Function of Desmosomes
+ The function of desmosomes is to adhere cells together.
‘+ They are found in high numbers in tissues that are subject to a lot of mechanical forces.
+ For example, many are found in the epidermis, which is the outer layer of skin, and the
myocardium, which is muscle tissue in the heart,
+ They are also found in between squamous epithelial cells, which form the lining of body parts
like the heart, blood vessels, air sacs of the lungs, and esophagus.
+ Numerous studies have characterized roles for desmosomal proteins in the regulation of
different signaling pathways.
Structure
Proteins from three main families coordinate to perform this function .
+ The junction has a dense cytoplasmic plaque composed of a complex of intracellular anchor
proteins, plakogiobin and desmoplakin that are responsible for connecting the cytoskeleton
intermediate filament to the transmembrane adhesion proteins.
+ These adhesion proteins, desmoglein and desmocollin like those at an adherens junction,
belong to the cadherin family.
+ They Interact through their extracellular domains to hold the adjacent plasma membranes
together.
+ Desmoglein and desmocollin are members of the cadherin family of adhesion proteins. Their
cytoplasmic tails bind plakoglobin (y-catenin), which in turn binds to desmoplakin. Desmoplakin
also binds to the sides of intermediate filaments, thereby tying the desmosome to these
filaments.dermogieig __deamocoltin
‘CYTOPLASM
plakogtobia
desmaplakin
wo Intermediate filaments
o-caterin Cosherins
Gap Junctions
+ Gap junetions in animal cells are like plasmodesmata in plant cells in that they are channels
between adjacent cells that allow for the transport of ions, nutrients, and other substances that
enable cells to communicate
‘+ This channel is big enough to allow small molecules such as inorganic fons, and other small
‘water soluble molecules (smaller than 1000kDa) to pass between the cells. However the channel
Is too small for proteins, nuclele acids or sugars to pass through.Plasma membranes
Intereettutar spac;
24 0M SECC Hy ampnilic channel
+ The membranes of two adjacent cells are separated by a uniform narrow gap of about 24 nm
+ maximal functional pore size for the connecting channels of about 1.5 nm
+ Gap junctions develop when a set of six proteins (called connexins) in the plasma membrane
arrange themselves in an elongated donut-like configuration called a connexon.
+ When the pores {“doughnut holes”) of connexons in adjacent animal cells align, a channel
between the two cells forms,
‘+ Gap junctions are particularly important in cardiac muscle: The electrical signal for the muscle to
contract Is passed efficiently through gap junctions, allowing the heart muscle cells to contract
in tandem.
Examples:The action potential in heart (cardiac) muscle flows from cell to cell through the heart
providing the rhythmic contraction of the heartbeat,
+ At some so-called electrical synapses in the brain, gap junctions permit the arrival of an action
potential at the synaptic terminals to be transmitted across to the postsynaptic cell without the
delay needed for release of a neurotransmitter,
+ As the time of birth approaches, gap junctions between the smooth muscle cells of the uterus
enable coordinated, powerful contractions to begin.‘Several inherited disorders of humans such as certain congenital heart defects and certain cases
of congenital deafness have been found to be caused by mutant genes encoding conn
s.
Plasmodesmata
Plasmodesmata (singular, plasmodesma ) are cytoplasmic channels thet pass through the cell
walls of adjacent cells,
Plasmodesmata are lined by plasma membrane and usually contain a dense central structure,
‘the desmotubule, derived from the smooth endoplasmic reticulum of the two cells,
Like gap junctions between animal cells, plasmodesmata serve as sites of cell-to-cell
communication, as substances pass through the annulus surrounding the desmotubule
plasmodesmata allow much larger molecules (up to SOkDa) to pass between cells,
plant viruses spread from one cell to anther through plasmodesmata
Plasmodesmata have been shown to transport proteins {inclucing transcription factors), short
interfering RNA, messenger RNA, viroids, and viral genomes from cell to cell
Plasmodesmata provide an easy route for the movement of ions, small molecules like sugars
and amino acids, and even macromolecules like RNA and proteins, between cells,
The larger molecules pass through with the aid of actin flaments,
smooth
endoplasmic
reticulum — desmotubule
cell walls
of adjacent
plant cells
ae cytosol
:
plasma membrane lining
plasmodesma, connecting
two adjacent cells