Cell Migration
Here we see a blood vessel lined with endothelial cells with a
normal flow of blood containing lymphocytes. We are going to follow one
particular lymphocyte to see how it migrates through the endothelium and out
into the surrounding tissues.
As the vessel widens into what is known as a venule, the blood flow
decreases and the lymphocytes slow down.
In the presence of an infection such as bacteria, an immune
response takes place and cytokines such as interleukin 1 and tumour necrosis
factor are released from tissue cells and resident macrophages. In addition,
peptides of the complement system cause mast cells to degranulate.
Cytokines such as interleukin 1 and tumour necrosis factor act on
the endothelium to induce adhesion molecules, while chemokines attach to the
endothelium to act as stimulators of migration.
The cytokines induce the expression of adhesion molecules on the surface of
endothelial cells complementary to those on the lymphocytes. Molecules on the
endothelial cells such as E-selectin interact with carbohydrate molecules on the
surface of the lymphocyte such as CD15.
These interactions cause the lymphocytes to slow down and roll
along the endothelium enabling them to sample other adhesion molecules which
have been induced onto its surface.
Amongst these are the chemokines which bind to the lymphocyte and trigger an
activation mechanism causing an influx of calcium ions into the cell and the
activation of another group of adhesion molecules onto its surface.
These adhesion molecule proteins called LFA1 and VLA4 interact with other
adhesion molecule proteins on the surface of the endothelium called ICAM1 and
VCAM1 respectively.
These latter molecules can also be induced onto the endothelial surface by
cytokines such as tumour necrosis factor thereby contributing to the overall
induction process. This multi-point adhesion onto the endothelium is called
pavementing.
Both the adhesion molecules on the lymphocyte and those on the endothelium are
each attached to their respective cell's cytoskeleton. This can be thought of as
a molecular motor which acts like a tractor enabling the lymphocyte to pull
itself down and out through the endothelium.
With a firm anchoring point to move against, the lymphocyte
proceeds to extend its pseudopod down into the intercellular junction between
the endothelial cells which retract as they are pushed down.
When the pseudopod makes contact with the basement membrane, the cell moves new
adhesion molecules down to its leading edge enabling it to form a new anchoring
point.
The lymphocyte now releases granules from this leading edge. These
contain destructive enzymes such as collagenase and elastase, which puncture the
basement membrane allowing the cell to squeeze down through the damaged area and
migrate out into the tissue. This migratory process is called diapedesis.
The damaged endothelium then seals over again and reforms.
The lymphocyte then migrates towards the site of infection under
chemical stimuli in a process known as chemotaxis.
Another feature of inflammation is an increase in capillary
permeability. Mediators from activated mast cells such as histamine;cauce the
endothelial cells to retract. This allows larger serum molecules such as
antibody and complement to leak into the inflamed area.
These mediators can also act on the smooth muscle in the
blood vessel wall, increasing the diameter of arterioles and producing enhanced
blood flow.
This causes increased supply of serum molecules and cells to
the inflammatory site.