Cardiovascular System: Myocardium and Heart

The cardiovascular system can be anatomically subdivided into the heart and the
blood vessels, and the latter category is further subdivided into different vessel
types.
Arteries are blood vessels which conduct blood away from the heart; veins are blood
vessels which conduct blood towards the heart. These two major categories are
broken down further, and there are subcategories of arteries and veins based on
size, construction of the vessel wall, etc. However, the direction of blood flow is the
criterion which separates arteries from veins, and which defines a vessel as being in
one or the other major class. You may have heard the hoary old chestnut that
"Arteries carry oxygenated blood, and veins carry deoxygenated blood." THIS IS
NOT TRUE. Some arteries carry deoxygenated, and some veins carry oxygenated
blood. The direction of flow is the only criterion for classification.
.
The Heart

Begin with slide 616, and start by

holding it up to the light. This is
an entire heart from some small
animal (most likely a rat) which
has been sectioned from the apex
to the cranial end of the atria.
After orienting yourself, place it

on the microscope under

low power. You will easily
be able to make out at
least two, and probably
three of the chambers; if
you have a favorable
section, you may be able
to see all four. The atria
and ventricles are
separated from each
other. Running from the
apex to the cranial end
you will see the septum
dividing the right and left
sides. There should be at least one of the heart valves in your section, connecting the
atria and ventricles. External to the heart proper, you should have a cross section of
one of the great vessels (the pulmonary artery) and some adipose connective tissue
as well.
Turn now to slide 66. This is the heart of yet another unfortunate rodent, sectioned
transversely, below the level of the atrioventricular junctions. This slide graphically
illustrates the thickness of the wall of the left ventricle and ventricular septum. The
cross section at right shows a "slug" of blood frozen in its passage from the left
atrium to the left ventricle, and the left A-V valve is open. On this slide you should
be able to make out most of the features identified in the previous slide, except those
which are out of the plane of the section.
While both atria and ventricles are composed of the same type of specialized muscle
cells, those of the atria tend to be less numerous, thinner, and more elongated than
those of the ventricles. The ventricles of the heart are more stoutly constructed than
the atria because they have more work to do. All the atria do is pass blood to the
ventricles below: the right ventricle sends blood to the lungs against the resistance of
the pulmonary circulation, and the the left ventricle has to deal with the entire
system circulation's resistance to flow. While the resistance of the lung capillaries to
blood flow is considerable, that of the entire peripheral circulation is much higher.
Myocardium
The heart is a large mass of muscle. Cardiac muscle is a variant form of striated muscle,
with distinct differences from the skeletal form; and some unique structures that make it
work properly, day in and day out, until death. The term "myocardium," (from Greek,
myos = muscle + kardio = heart) is specifically the mass of muscle that comprises the
bulk of the organ.
On slides 616 and 66 at low power, you can easily observe the pattern of the muscle
cells of the myocardium. Myocardial cells (cardiac myocytes) are much smaller than

the myofibers of skeletal muscle, and they don't form long cylindrical structures the
way myofibers do. While skeletal myofibers can in some cases be meters in length,
it's a rare cardiac myocyte that exceeds 100 m in length, and perhaps 30-50 m
wide. The shape of the myocyte is pretty irregular, with stumpy projections coming
off of it where it contacts other cells. Myocytes also differ from skeletal myofibers in
that cardiac myocytes are mononuclear and not syncytial. One nucleus per myocyte
is the rule, not hundreds as in a skeletal myofiber, and that one nucleus is centrally
located.
Here's scanning
electron
microscope
image of a
single cardiac
myocyte, which
has been
mechanically
separated from
the mass of the
myocardium.
This beautiful
image (at about
5000x) shows
the irregular
shape of the
myocyte quite
well, and the
points at which
this cell is in contact with others via the intercalated discs are indicated by arrows.
Also visible as wispy strands of material on the surface are fine collagen fibrils of
the intercellular collagen network. Just as in skeletal muscle, the ICN transmits
force throughout the mass, and the hierarchical arrangement of endomysiumperimysium-epimysium applies.
Note that the surface of the myocyte appears to be ridged or grooved. These ridges
are the sites of Z-lines. They mark the ends of the sarcomeres of this cell's
contractile apparatus. The sarcomeres are slightly contracted in this specimen and
so they appear as ridges on the
surface. The T-tubules aren't
visible.
In the image at right, a whole
mount stained with toluidine
blue at about 1000x, two
myocytes have come through
the disaggregation procedure

but remain connected to each other: one is more heavily stained than its partner, so
that the boundary where the two are joined (an intercalated disk, see below) is clear.
As is obvious in this image, each cell has a single centrally located nucleus. Unlike
skeletal muscle, neither myocardium nor the cells of which it's composed are
syncytial. The cells are physically isolated from each other, a fact which was in
dispute before the electron microscope settled the matter once and for all about
1950. (Nevertheless, thanks to the Law Of Persistence Of Erroneous Information,
you'll still find the statement in texts that "the heart is functionally syncytial,"
something that sets my teeth on edge.) Though there is undoubtedly constant
communication and coordination between heart muscle cells, they are structurally
distinct entities, and the term "syncytium" is inappropriate in this context.
Thanks to the difference of cellular architecture, the histological appearance of
myocardium is very different from that of skeletal muscle. Rather than forming neat
bundles of parallel fibers with well-defined striations, myocardium forms an
anastomosing network with a sort of "spongy" appearance. The contacts between
myocytes are such that a branching network with blood vessels in the spaces between
them is the result.
You will see some areas where the myocardial cells are oriented end-to-end in long
rows parallel to the plane of the section (i.e., they are cut longitudinally) and others
where they run at right angles (and thus are cut transversely). There will also be
areas where the orientation with respect to the plane of the section is less well
defined; these are oblique sections. The myocardial muscle bundles are oriented in
such a way as to make most efficient use of the force of contraction.

These two images give a pretty good idea of the appearance of myocardium in the
light microscope. The one at left is stained with toluidine blue at about 200x; the one
at right with H&E at roughly 400x.
Myocardium has a "stringy" look compared to skeletal muscle. While skeletal
muscle cells are very large and lie next to each other in parallel bundles, the smaller
cardiac muscle cells are butted together at their ends, and irregularly shaped,

numerous blood vessels between them. The effect is to create an anastomosing

network of fibers rather than a solid phalanx of muscle fascicles. The single nucleus
of each myocardial cell is quite clearly visible in these images, especially on the
right.

This drawing will clarify the architecture of the tissue. Note the branching, and
compare it to the actual specimens shown in the longitudinal view. At lower right in
the drawing, the cells are shown in cross section. Compare the drawing to the actual
specimen shown at right, and to the sections of smooth muscle in Exercise 10: at first
glance this field could be confused with smooth muscle, but they can be told apart
fairly easily. An important difference between the appearance of this tissue
compared to smooth muscle is that in cardiac muscle almost all the cell profiles are
approximately the same size, which isn't true of smooth muscle. Furthermore, most
of the cell profiles will have a nucleus inside. Cardiac myocytes are so short and the

nucleus takes up such a proportionally greater percentage of their length (compared

to smooth muscle cells) that the chances of intersecting a nucleus with the plane of
the section is pretty good. The individual cells are clearly defined by their CT
envelope (the endomysium) and most of them show a nuclear profile.
The smaller, denser nuclei outside the cells are fibroblasts that make and maintain
the intercellular collagen network. There's a blood vessel crossing the field about
one quarter of the way down from the top edge. Cardiac muscle is even better
served by the circulatory system than skeletal muscle is. Unlike skeletal muscle,
cardiac muscle can't incur an "oxygen debt," because it doesn't have the lactic acid
pathway to generate ATP when oxygen levels are low. Consequently it's quite prone
to anoxia, and the capillary beds are extensive to minimize the possibility of it
happening.
Intercalated Disks
The most prominent feature of myocardium, and the one that's absolutely
diagnostic for it, is the intercalated disk. The ID is a specialized cell-to-cell
adhesion/communications site. It demarcates the beginning of one myocyte and the
end of the next, and information is pass across it from cell to cell. These structures
are found only in cardiac muscle.
The ID's are vital to normal myocyte function and understanding how they are put
together is important. In these special areas there's a considerable degree of
interdigitation of myocyte plasma membrane, but there is no actual fusion of
cytoplasm. The specialized apparatus of the ID allows myocytes to act as if they
were a true syncytium there is in fact no actual cytoplasmic continuity; these are
distinct and individual cells.
The true nature of the ID wasn't fully understood until the transmission electron
microscope was available for their study. This instrument revealed that ID's are
physically held together by large numbers of desmosomes, with gap junctions
between the myocytes forming a region that permits electrical communication in the
form of ion fluxes. This flux maintains the coordination of the waves of contraction
between one cell and the next.

Intercalated discs are readily seen on slide 525, in regions where the myocardium is
cut longitudinally (they won't be visible in cross sections). Since the ID's are located
at the ends of the cell, and since that's where the first and last sarcomeres of any
given myofibril are, you can think of the ID as a sort of thickened "terminal Z-line."
ID's are just barely at the limit of visibility in a light microscope, and it may help to
defocus your substage condenser a bit. They'll be seen as fuzzy lines running across
the width of a row of cells. The presence of ID's is absolutely diagnostic for cardiac
muscle; no other tissue has such structures.
Laterally there are no points of fusion or adherence between cells that resemble the
ID's. They're only at the ends of the cells. The ID acts as a communication
site as well as an adherence structure between two myocytes. Gap junctions
between the two myocytes take care of the communication: the ID is a site of
lowered electrical resistance, so that passage of ions through the gap
junctions into the intercellular space keeps adjacent cells "informed" about
what's happening. The coordination of the heartbeat and the transmission of
the contraction signal from one cell to the next is dependent on the integrity
and proper functioning of the ID's gap junctions. Cell-to-cell adherence is
maintained here (as at other sites) by numerous desmosomes. These are
scattered along the length of each ID.
Endocardium and Epicardium

Examine the lining of the chambers on slide 616. The entire cardiovascular system,
including all chambers of the heart and all blood vessels, is lined with a simple
squamous epithelial covering. This lining is among the most metabolically active
tissues in the body. In blood vessels, this is (by convention) called "endothelium,"
but in the heart the special term endocardium is used instead. (There isn't a nickel's
worth of difference between them, but the terminology is so entrenched it could
never be eradicated.) In any
event, "endocardium" is an apt
name because it's literally
"inside the heart." The
endocardium covers the valve
cusps, too, as you can easily
verify on this slide. If you
traverse the thickness of the
myocardium and examine the
outer surface of the wall, you
will see a similar (though less
well defined) layer of simple
squamous epithelium. Again, by
convention, this has a special
name; it's the epicardium, which
translates as "upon the heart."
Anatomically, this is the visceral
layer of the pericardial sac.

The epicardium is the inner portion of the

CT envelope that surrounds the heart. It
corresponds to the peritoneum of the
abdominal cavity in origin (from the
mesoderm lining the embryonic coelom)
and function (to allow the heart to move
freely in its cavity without adhesion to
surrounding structures). Here you see it as
a thin serous membrane overlying the
outer surface of the organ. A stain for
connective tissue would reveal a very
delicate sub-epicardial CT layer, mainly
reticular fibers.
Purkinje Fibers
Slide 582 is another chunk of myocardium. Most of the features of myocardium are
visible, but this section also shows Purkinje fibers (Johannes Evangelista von
Purkinje, 1787-1869, a Czech anatomist, physiologist and microscopist). These fibers
are part of the conducting system of the heart.

The conducting system transmits the signal to contract from its origin in the
"pacemaker" to the rest of the myocardium. The Purkinje fibers are not neurons.
They're specialized muscle cells, and they look like it. They can be distinguished
from the myocardium by their size (they're much larger than regular myocytes) and
by their
staining
(they tend
to be more
lightly
stained
than true

myocardium). With the PAS stain they're strongly positive, since they contain large
amounts of glycogen.
Purkinje fibers are not contractile, at least not to any significant extent. They
contribute nothing to the force generation of myocardium. The cells of the Purkinje
fibers are organized into several tracts that lead away from the atrioventricular
node and form nerve-like structures. There's a bundle along both sides of the
septum, one distributing to the right ventricle and one to the left ventricle. The
Purkinje fibers are larger and less well stained than the ordinary contractile
myocardium because of the relatively protein-poor cytoplasm, which doesn't take up
the eosin stain very well. Hence the fibers look pale and washed out in comparison
to the protein-packed myocytes. Purkinje fibers do have some of the other features
of cardiac myocytes, including a centrally-located single nucleus, and even
intercalated disks.
But despite their strictly conductive function, they are muscle cell derivatives, so
you're quite likely to see striations and other indications of "contractility" if you
examine them at higher power. There should be two areas of Purkinje fibers in your
slide, one at each edge of the section. In this slide also note the ramifications of the
muscle, and the intercalated discs.
The Purkinje fibers ramify into the mass of ventricular myocardium, and from their
termini the signal is spread through the ID system. Recall that all muscle tissue is
"excitable" and this property can be used to carry information. That's what's

happened here. What is a secondary function in contractile myocytes has been

transmuted into the main role of these not-quite-muscle cells. The fibers run
through the sub-endothelial connective tissue; they can't generate any force because
internally they have only a few disorganized actin and myosin fibrils, nothing like
the very regular arrays of sarcomeres in the contractile cells.
Slide 96, shown at left, is
interesting because it contains the
atrioventricular node, the
beginning of the Purkinje fiber
conduction apparatus.
This image is taken with the atrial
region at the top of the field. At
the upper left is the base of one of
the great vessels leading from the
heart (probably the aorta); and
below that a triangular-shaped
region, the trigona fibrosa. The
trigona fibrosa is part of the socalled "cardiac skeleton," the
dense fibrous CT to which the
intercellular collagen network is
ultimately anchored. If you look
at this structure at high
magnification it's very similar in
appearance to cartilage. The
trigona fibrosa lies between the
atrial and ventricular parts of the
organ and the fibrous rings of the
great vessels are attached to it; so
is the top level of the ICN. When
force is exerted by contraction of
myocardium, this is the "zero
point" against which the force is exerted. When contraction occurs, the different
directions of muscle tracts impart a compressive, twisting motion to the heart
grossly (you will be able to see this in surgery). The motion has been compared to
"wringing out" a wet cloth, and it serves the same function: expulsion of the
maximum amount of blood per stroke. Contraction of the myocardial cells leads to
reduction in volume of the heart chambers, and consequently to ejection of all (or
almost all) of the blood therein. Impairment of this action results in decreased
efficiency of pumping and eventual failure.
Alongside the trigona, and lyingas its name impliesbetween the atrium and the
ventricle, is the A-V node. It looks like a bit of atrial myocardium, splonked up
against the denser and more massive ventricular type. (This slide also gives you a

good comparison of the two types of myocardium side by side). This isn't the
"pacemaker," (that's the sino-atrial or S-A node) but it receives signals from it and
transmits them via the conducting fibers to the mass of ventricular myocardium.
Histologically it looks like the rest of the atrial myocardium, and in the absence of
the landmarks used here, would be very difficult to identify in a section.
Valves of the Heart
Slide 102 is of a valve cusp. Look at it at low magnification and at high
magnification and examine its construction. The cusp of a valve has a core of CT,
much of it composed of elastic fibers. There

may be some smooth muscle worked into it as well. The outer surfaces, exposed to
the blood flow, are covered with the epithelium of the endocardial lining, like the
rest of the system. Note that there's an excursion of myocardium into the base of the
valve cusp; this terminates before the end of the cusp. It's continuous with the
myocardium of the ventricle.
Blood Supply
I hope it hasn't escaped your notice that the entire mass of the myocardium is shot
through with blood vessels of varying sizes and shapes, and that there's an extensive
degree of vascularization. In fact, the heart pumps blood to itself before it sends any
to other parts of the body. The first branches off the aorta are the coronary arteries
of the myocardial circulation. After you have completed the section on blood vessels
you may want to come back and classify some of these.
Arteries
Let us now leave the pumping station and examine some of the pipes: the arteries
and veins. Remember the definition: arteries pump blood away from the heart;
veins carry blood towards the heart. No other criteria for classification of a vessel as

one or the other exists. Capillaries are those small vessels within the tissues from
which oxygen transfer takes place.
Arteries may be subclassified by type. Conducting or elastic arteries are large ones,
with very strong and relatively elastic walls, whose function is to "conduct" the bulk
of the blood to regions of the body where it's to be distributed. Examples include the
aorta, subclavian, and pulmonary arteries. Once the blood has reached the region of
distributionsay, the limbsit will be handled by smaller (but still fairly large)
distributing or muscular arteries, which send it to sub-regions. As the distribution
area gets more and more limited the arteries become smaller. In very local areas you
will see small arterioles, essentially mini-arteries with a wall considerably less
muscular than the larger ones "upstream."
Elastic or Conducting Arteries
Elastic arteries are constructed like fire hoses. Rightly so, because they have the
same function: to carry a stream of liquid under high pressure. Hence they're
designed to minimize internal friction and flow resistance; and to maximize the
strength of the wall.
The lumen is lined with a thin squamous epithelial layer (the tunica intima) which
may be likened to the rubber bore of the hose; like the rubber, it offers a smooth and
unimpeded passage for the flow of blood. The tunica media is a region of elastic and
collagen fibers. Elastic arteries (the aorta most of all) must withstand an enormous
head of pressure to pump against the peripheral systemic resistance: consequently
the wall is heavily reinforced to prevent bursting, just as the wall of a fire hose has
reinforcing cords in it. The elastic fibers allow some stretching and "springiness" in
response to the pressure, and the collagen fibers limit the degree of stretch
permitted. (In some disease or deficient nutritional statesfor example lathyrism, a
copper deficiency caused by certain plantsthe wall may be weakened, resulting in
an aneurysm which may leak, or burst with fatal effects.
To complete the analogy, the collagenous tunica adventitia of the aorta is the fabric
covering on the outside of the hose. Just as the fireman needs a firm grip to control
the hose, so must elastic arteries be anchored down to the surrounding structures, to
prevent them from moving around as pressure varies internally. The tunica
adventitia of conducting arteries is scanty, and collagenous in nature.
Slide 30 is a fine example of an elastic or conducting artery. It is, in fact, a section of
the aorta. The section on this slide is stained with the Verhoeff's stain for elastic
fibers. It renders these fibers black, and you will see that the wall of this vessel is
shot through with elastic fibers. The spaces between the elastic fibers are mostly
occupied with collagen, and some small amount of smooth muscle (see below). The
amount of elastic fiber infiltration is so great that no internal or external elastic
laminae can be identified.

This is a section of the aorta, a nice example of an elastic artery. The elastic fibers
normally aren't easily seen in an H&E preparation like this one, but in this case, the
reinforcement is so heavy that the concentric layers of elastic CT show up as distinct
iridescent rings, some of which are marked by arrows. There's a large clot of blood
in the field. The bore of this vessellike the rest of the cardiovascular systemis
lined with a simple squamous epithelium.
The innermost of the numerous elastic layers in this artery has its own name: the
internal elastic lamina. It's the one right up against the tunica intima. The internal
elastic lamina is much more easily seen in muscular arteries (see below) than in the
elastic ones, though. In an elastic artery like this one it gets lost in the "background"
of dozens of similar layers.
As is true of most elastic arteries, the tunica adventitia on this one is a relatively
small contributor to the wall's thickness.
There's more than elastic fibers in the wall of this type of artery. Between the elastic
layers there is a considerable amount of collagen and some smooth muscle,
permitting the artery to expand under pressure and recoil to original diameter when
the pressure drops again. Collagenous components in the wall prevent overexpansion and resist bursting of the vessel.

Using stains specific for elastic fibers reveals how extensive the reinforcement of the
wall can be. Here are two examples, again from the aorta. The one at left has been
stained with a combination of the Verhoeff and the Van Gieson stain; at about 400x,
the extent of the elastic component (in black) and the collagenous reinforcement
between the elastic fibers is easily visible. This field is from the outer edge of the
tunica media: the almost-completely red area at the bottom is the tunica adventitia.
At right, the Verhoeff stain has been used and at about 200x it clearly show elastic
fibers but not the collagen or smooth muscle, which is unstained by this method.
Very large elastic arteries have their own internal blood circulation system and
nervous supply. They have to: the fibroblasts and other cells that keep the wall in

good shape are so far from the blood supply that diffusion won't serve their needs,
and the density of the wall and its fibrillar components also impede diffusion. The
smooth muscle in the wall is innervated so that the CNS can control blood pressure
and initiate contractions when needed. The vasa vasorum (and in the case of nerve
fibers, the nervi vasorum) i.e., the "vessels of the vessels" and the "nerves of the
vessels" are a constant feature of big vessels.
Muscular Arteries
Once you get
the blood out
to the major
regions of
the body,
there's a
transition in
the structure
of the
arterial wall.
The
proportion
of elastic
fibers
decreases,
and the
proportion
of smooth
muscle
increases.
Some elastic
fibers and
collagen
fibers will
always be
present, but
eventually
the great
bulk of the tunica media will be smooth muscle, and at that point we're dealing with
muscular or distributing arteries, whose function is to "distribute" blood supply to
their regions of responsibility, such as a limb. The artery on slide 103, at right, is a
dandy example of what a muscular or distributing artery should look like. This is
the femoral artery; the femoral vein (see below) and the femoral nerve are there as
well. There may be a branch point off this artery on your slide.
Note that the wall of the artery is mostly tunica media, and that this tunic is almost
entirely smooth muscle. Elastic CT is present, to be sure, but not nearly to the extent

that it is in the elastic arteries. There is also a noticeable internal elastic lamina,
stained bright pink, and a less well defined external elastic lamina, though at the low
magnification of this image it's hard to make out. The endothelium of the tunica
intima is easily visible. The tunica adventitia grades off into the surrounding
connective tissue, but is fairly sharply defined. The internal elastic lamina is just
barely visible in this image. It's the undulating pink line immediately below the
lining endothelium. It marks the innermost limit of the muscular tunica media. The
external elastic lamina is less regular and can't be made out at all. Note the
prominent collagenous tunica adventitia here. In muscular arteries the tunica
adventitia is often most of the wall's overall thickness. The external CT investment
anchors this artery to the surrounding CT.

The internal elastic lamina is most easily seen in smaller muscular arteries. In these
it usually stands out as a bright pink undulating band just below the lining epithelial
cells and their supporting CT. In life, the artery is always under some pressure, but
when death occurs the tonus of the wall causes a partial collapse, so that the
undulation is something of an artifact.

In the example at left, the IEL is indicated; this example also clearly shows the
separation of the lining epithelium from the IEL by the sub-endothelial CT. A few
layers of smooth muscle constitute the tunica media in this small artery.
The smooth
muscle of the wall
of distributing
arteries makes
them very
extensible, and
also provides for a
counter force to be
exerted against
the pressure of
filling. As the
vessel expands the
smooth muscle
cells are stretched:
in reaction to this
they begin to
contract. The
peak of their
contraction comes
at about the point where systole ends and diastole begins; thus the contraction of the
arterial walls dampens out the pulsations of the flow to provide a more or less steady
supply of blood at normal pressure into the capillary beds. Nervous input can also
control this to some extent, independent of the mechanical force of stretching. As
distance from the heart increases, the force required to dampen the oscillations is
less, and smaller arteries can handle it; the interval between peaks also lengthens
and there is a much more uniform flow rate.
It's
useful to
put the
two
major
arterial
types
side by
side,
stained
to reveal
the wall

components. On slide 30, this has been done using the Verhoeff stain. A small
muscular artery is near the large elastic one, to make the point that as the artery
size decreases, the proportion of elastic fibers in the wall decreases as well. The wall
of the small artery has almost no elastic tissue in it, but the the internal elastic
lamina, however, is very clearly defined against the background of the muscular
tunica media.
It's possible to combine several staining methods to show all the major components
of the wall, including the smooth muscle. In the image below, the Verhoeff and
Masson's stains have been combined. Elastic tissue is stained black; smooth muscle
is red; and collagen is green. The tunica media (TM) is almost entirely muscle, with
a few minor streaks of green collagen in it. The inner elastic lamina (IEL) stands out
prominently; and the outer elastic lamina (OEL) demarcating the end of the tunica
media is also easily visible. The tunica adventitia (TA) is a mixture of green collagen
fibers and black elastic fibers interwoven with each other to provide strength and
resilience.
Slide 8 demonstrates the brachial artery and vein. The artery has the typical
structure of a distributing artery, and it has a very nice tunica intima and tunica
media. The vein shows the typical structure of tunica intima, scanty tunica media,
and a thick CT tunica adventitia. There are also valves present.
Arteries In Longitudinal Section
Since arteries are pretty common in almost any microscope slide you'll look at, it's
important to be able to recognize them when they aren't cut in cross section, as all
the previous examples have been.

Here are two examples from tissue sections. The left section is from slide 34; the
right from slide 148. Since the smooth muscle in the wall of an artery is oriented
with the long axis of the smooth muscle cells around the long axis of the vessel
proper, in cross sections, the muscle cells are cut in longitudinal view, but in long
sections of the vessels, as here, the cells are cut in cross section. Compare these
profiles (both at about 400x) to the sections of smooth muscle in Exercise 10, and
you'll be able to make out the boundaries of the smooth muscle easily.
Aneurysms
Sometimes the wall of an arteryespecially a big one like the aorta, which is
subjected to all or nearly all the pressure the heart can generateis weakened by
disease, malnutrition, age, or some other affliction. Under a good head of pressure,

blood begins to dissect the layers of reinforcing material, separating them and
causing the side of the artery to bulge, exactly as a garden hose does when its wall is
weakened. This bulge is an aneurysm. Chronic high blood pressure makes this more
likely, though an aneurysm can occur even when systemic blood pressure is lower
than normal. Needless to say, an aneurysm that bursts will really spoil your plans
for the weekend. If it's a big one that blows out, such as the aorta, death will be
rapid.
Veins
Veins are
those
vessels
leading
blood back
towards
the heart.
As a rule,
they have
much
thinner
walls than
arteries
do, though
in cross
sectional
area
they're
usually
larger
than the
corresponding artery, because they have to carry the same volume of blood at a
lower pressure. Since veins are on the post-capillary side of the circulatory loop,
operating at much lower pressures, there's less need for burst resistance. Thin walls
are also important because much of the pressure that drives blood through veins is
generated not by the heart, but by contraction of the muscles in the region of the
vein. This "squishes" the blood back through the vein. Because they have low
pressures, some veins have venous valves in them to prevent back flow. This is
especially true of medium sized veins in the extremities, as they have to lift blood
against gravity.
(If an animal is held totally immobile for a long period of time, the return of the
blood to the heart is diminished, because contraction of the muscles of the limbs no
longer pushes blood back to the heart. Since the heart can only put out what it takes
in, the net result of decreased venous return is decreased arterial output. At some
point the output declines to the point where the brain is no longer receiving

sufficient blood, and the animal loses consciousness. This happens surprisingly
often. Soldiers on parade, when forced to remain at the position of "attention" for
long periods, will from time to time keel over and quite literally "drop out" of the
ranks as they
faint.)
Venous Valves
Veins of a
certain size
usually have
valves to
prevent back
flow. Veins of
this type are
usually found
in the
extremities, the
valves allowing
blood to move
back towards
the heart with
less effort.
Venous
pressures are
so low, and the
propelling
force has to
fight against
not only the resistance of the vessels but the relentless pull of gravity, that valves in
the vein allow the blood to be pushed up above and when the valve closes without
running back down when pressure slacks off.
The venous valve has a great deal of structural similarity to the valves of the heart.
There's a CT core with epithelium on both sides. Two valve flaps meet in the center
of the vessel. You can find such valves on slide 8, and when you do, note especially
the direction of the valve: it's designed to permit blood to flow in one direction only.
Pooling of blood occurs on the superior side of the valve flaps. In animals with long
lives and vertical postures, the continual stress that the weight of this blood imposes
sometimes causes outpouching of the vein. If there is a weakening of the wall on the
downstream side of the valve, the blood will push it out into a small bubble
analogous to the aneurysm in an artery. The result is what we call "varicose veins"
that many peopleespecially those who work on their feetdevelop in later life.
This image from slide 8 shows a fairly large vein, cut more or less in cross section.
The flaps on both sides of one of its valves are visible.

Capillaries
Capillaries are small vessels, whose walls are thin enough to allow the diffusion of
nutrients, oxygen and carbon dioxide across them. They are "where the action is" in
gas and nutrient exchange: with a few exceptions, no cell of the body is very far
from one, because access to the blood is an absolute requirement: cell death would
result from anoxia or the loss of nutrient and waste transport.

Capillaries are by far the most numerous class of vessels, though they're so small
they can only be appreciated in microscopic sections. There are two types: closed or
continuous capillaries, and fenestrated capillaries.
The closed type is found in locations where rapid "bulk" transfer of materials
between the blood and the tissue it served isn't needed, such as in muscles. Closed
capillaries can move material in and out using a process of sequential endocytosis
and exocytosis, as well as simple diffusion for small molecules.
Fenestrated capillaries, which have actual pores in their walls, are located wherever
immediate movement of materials is a functional necessity, such as in endocrine
organs and in the kidney. They don't demonstrate the "transcytosis" activity of the
closed type, because bulk movement of ions, hormones, nutrients, etc. through the
pores is ample

Here's a scanning
electron micrograph of
a capillary running
alongside muscle
fibers. This is a closed
capillary. This striking
image shows the
approximate size of a
capillary very well:
they're just about the
same size as the
erythrocytes that flow
through them, maybe
10 m in luminal
diameter. In this
picture you can see an
erythrocyte peeking
out at the broken end,
and the anchoring
fibrils of connective
tissue that attach it to
the surrounding muscle mass (RF).
Compare this to the diagram above: the wall is made of very thin squamous cells
which are sealed together at the edges by desmosomes; in other words, there's only a
tunica intima, and not much of that beyond the lining epithelium and the basement
membrane it rests on. In a section it's normal to see parts of several of the mural
cells in the same plane, since there's quite a bit of interdigitation between the
irregular borders of adjacent cells.
You may also see a second cell type, the pericyte. Technically this isn't part of the
capillary wall. Pericytes "embrace" the wall but never make contact with the blood,
as the mural cells do; they're believed to have some sort of contractile function.
They're so closely associated with the mural epithelium they share a basement
membrane with it.

Despite
their small
size,
capillaries
are so
numerous
that
they're
easy to
find. They
often will
have blood
cells in
them,
which also
makes
them easy
to spot.
This one is
from
skeletal
muscle on slide 14. Skeletal muscle has an abundant blood supply, and this capillary
is lying between two of the large muscle cells. A couple of erythrocytes are visible,
and the bore diameter of the capillary is just large enough to accommodate them
(double arrow). A white blood cell (probably a neutrophil) is visible in the left end of
the field. As it happens there are no capillary cell nuclei nor pericyte nuclei visible,
they're out of the plane of the section; but the wall itself can be see. The cytoplasm
of the mural cells is very scanty, and hence the precise limits of the cell and the CT
around it are hard to make out.
The capillary at right is
from white fat. It's cut in
cross section. The plane of
section has passed through
the nucleus of the mural
cell, and the lumen is filled
with the cytoplasm of an
erythrocyte.
All blood vessels, of
whatever type, are derived
from embryonic
mesoderm, the only one of
the three layers of the
embryo that has angiogenic
potential. It shouldn't

therefore be too surprising to find that connective tissues, which are also of
mesodermal origin, are usually well supplied with blood vessels. That's the case
here: white fat is a connective tissue, and the fibrous CT that separates each fat cell
(the clear spaces in this image) from the others is a second type of CT. This
capillary's function is to move the components of the fat stored in it into and out of
the fat cell, and to serve the needs of this tissue for nutrient and waste transport. In
young animals, which haven't had time to accumulate the "wear and tear" pigment
lipofuscin in their fat cells (see Exercises 2 and 3 for a discussion of lipofuscin) the
presence of so many capillaries give the fat a pinkish-white color. In albinos, animals
that lack melanin, it's the blood circulating in capillaries of the eye that's the source
of its pink color.
A fenestrated capillary has "holes"
in it; the word comes from fenestra,
the Latin for "window." These holes
are actually pores, places where the
plasma membrane is perforated, to
allow for the movement of bulk
materials from the capillary lumen
to the exterior, and vice versa. This
type of capillary is found in places
where the rapid movement of
materials is vital to function, such as
in endocrine organs, where
movement of hormones into the
blood is carried out. Fenestrated
capillaries are also found in the
kidney, again a place where quick
transit between two compartments is
the design criterion.
It's not possible to see the
fenestrations in a light microscope
preparation. To appreciate the
nature of these pores, an electron microscopic image is required, because they're
below the level of the light microscope's resolution. Such an image is provided here,
at about 80,000 diameters. This example is from the kidney, but similar capillary
profiles could be found anywhere fast movement of large molecules is important to
normal function. The capillaries of the glomerulus (the tuft of blood vessels that fills
the renal corpuscle) use hydrostatic pressure from the arterial supply to filter blood
plasma through the pores in the formation of urine (for details, see Exercise 23). As
this image makes clear, the fenestrations are actual openings in the capillary wall.
The arrows show the direction of flow of materials, driven by the higher pressure in
the capillary lumen than in Bowman's space. Remember, the image shown here is
two-dimensional, but the cell isn't. The "breaks" indicated are really more or less

circular perforations in the plasma membrane, and they have depth; they lie on both
sides of the plane of the section.
Sinusoids
There's one
more category
of blood vessels
to be dealt
with. These are
somewhat like
capillaries, and
might be
considered as a
sub-set.
Sinusoids are a
form of large,
irregular
capillary-type
vessel; they
have the thin,
intima-only
wall
construction of
a capillary and
may be
fenestrated.
Sinusoids are
found where
slow flow,
intimate contact between blood and tissue, and rapid exchange of materials are
required. You'll find them on slide 123 in the liver, between plates of hepatic cells.
Sinusoids are
flattened and
irregular in shape,
as the image at
right makes clear.
This scanning EM
picture is a plastic
cast of the
sinusoids in the
placenta of a goat,
another place
where slow flow
and efficient

transfer of materials is important to normal function. In this method the blood

spaces are filled with plastic; the plastic is allowed to harden, and the tissue digested
away to leave an accurate impression of the shape of the filled space.