Cardiovascular System From Veterinary Histology PR

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    Cardiovascular System: Myocardium and Heart

    The cardiovascular system can be anatomically subdivided into the heartand 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 ma!or categories are

    bro"en down further, and there are subcategories of arteries and veins based on

    si#e, 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 ma!or class. $ou may have heard the hoary old chestnut that

    %Arteries carry o&ygenated blood, and veins carry deo&ygenated blood.% TH'S 'S

    ()T T*+. Some arteries carry deo&ygenated, and some veins carry o&ygenated

    blood. The direction of flow is the only criterion for classification.

    .

    The Heart

    -egin with slide /, and start by

    holding it up to the light. This is

    an entire heart from some small

    animal 0most li"ely a rat1 which

    has been sectioned from the ape&

    to the cranial end of the atria.

    After orienting yourself, place it

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    on the microscope under

    low power. $ou will easily

    be able to ma"e out at

    least two, and probably

    three of the chambers if

    you have a favorablesection, you may be able

    to see all four. The atria

    and ventricles are

    separated from each

    other. *unning from the

    ape& 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. &ternal to the heart proper, you should have a cross section of

    one of the great vessels 0the pulmonary artery1 and some adipose connective tissueas well.

    Turn now to slide . This is the heart of yet another unfortunate rodent, sectioned

    transversely, below the level of the atrioventricular !unctions. This slide graphically

    illustrates the thic"ness of the wall of the left ventricle and ventricular septum. The

    cross section at right shows a %slug% of blood fro#en in its passage from the left

    atrium to the left ventricle, and the left A23 valve is open. )n this slide you should

    be able to ma"e out most of the features identified in the previous slide, e&cept those

    which are out of the plane of the section.

    4hile both atria and ventricles are composed of the same type of speciali#ed musclecells, 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 wor" 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 circulation5s resistance to flow. 4hile 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#ulk of the organ.

    )n slides / and at low power, you can easily observe the pattern of the muscle

    cells of the myocardium. Myocardial cells 0cardiac myocytes1 are much smaller than

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    the myofibers of s"eletal muscle, and they don5t form long cylindrical structures the

    way myofibers do.4hile s"eletal myofibers can in some cases be metersin length,

    it5s a rare cardiac myocyte that e&ceeds /66 7m in length, and perhaps 86296 7m

    wide. The shape of the myocyte is pretty irregular, with stumpy pro!ections coming

    off of it where it contacts other cells. Myocytes also differ from s"eletal myofibers in

    that cardiac myocytes are mononuclearand not syncytial.)ne nucleus per myocyteis the rule, not hundreds as in a s"eletal myofiber, and that one nucleus is centrally

    located.

    Here5s scanning

    electron

    microscope

    image of a

    single cardiac

    myocyte, which

    has been

    mechanicallyseparated from

    the mass of the

    myocardium.

    This beautiful

    image 0at about

    9666&1 shows

    the irregular

    shape of the

    myocyte uite

    well, and the

    points at whichthis 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 networ". ;ust as in s"eletal muscle, the 'C( transmits

    force throughout the mass, and the hierarchical arrangement of endomysium2

    perimysium2epimysium applies.

    (ote that the surface of the myocyte appears to be ridged or grooved. These ridges

    are the sites of

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    but remain connected to each other: one is more heavily stained than its partner, so

    that the boundary where the two are !oined 0an intercalated dis", see below1 is clear.

    As is obvious in this image, each cell has a single centrally located nucleus. +nli"e

    s"eletal muscle, neither myocardium nor the cells of which it5s 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/=96. 0(evertheless, than"s to the >aw )f ?ersistence )f rroneous 'nformation,

    you5ll stillfind the statement in te&ts that %the heart is functionally syncytial,%

    something that sets my teeth on edge.1 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 conte&t.

    Than"s to the difference of cellular architecture, the histological appearance of

    myocardium is very different from that of s"eletal muscle.$ather than forming neat#undles of parallel fi#ers with well%defined striations, myocardium forms an

    anastomosing network with a sort of "spongy" appearance. The contacts #etween

    myocytes are such that a #ranching network with #lood vessels in the spaces #etweenthem is the result.

    $ou will see some areas where the myocardial cells are oriented end2to2end in long

    rows parallel to the plane of the section 0i.e., they are cut longitudinally1 and others

    where they run at right angles 0and thus are cut transversely1. There will also be

    areas where the orientation with respect to the plane of the section is less well

    defined these are obliue sections. The myocardial muscle bundles are oriented in

    such a way as to ma"e 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 @66& the one

    at right with H at roughly B66&.

    Myocardium has a %stringy% loo" compared to s"eletal muscle. 4hile s"eletal

    muscle cells are very large and lie ne&t to each other in parallel bundles, the smaller

    cardiac muscle cells are butted together at their ends, and irregularly shaped,

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

    networ" of fibers rather than a solid phalan& of muscle fascicles. The single nucleus

    of each myocardial cell is uite clearly visible in these images, especially on the

    right.

    This drawing will clarify the architecture of the tissue. (ote 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 &ercise /6: 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

    appro&imately the same si#e, which isn5t true of smooth muscle. urthermore, most

    of the cell profiles will have a nucleus inside. Cardiac myocytes are so short and the

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    nucleus ta"es up such a proportionally greater percentage of their length 0compared

    to smooth muscle cells1 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 0the endomysium1 and most of them show a nuclear profile.

    The smaller, denser nuclei outside the cells are fibroblasts that ma"e and maintainthe intercellular collagen networ". There5s a blood vessel crossing the field about

    one uarter of the way down from the top edge. Cardiac muscle is even better

    served by the circulatory system than s"eletal muscle is. +nli"e s"eletal muscle,

    cardiac muscle can5t incur an %o&ygen debt,% because it doesn5t have the lactic acid

    pathway to generate AT? when o&ygen levels are low. Conseuently it5s uite prone

    to ano&ia, and the capillary beds are e&tensive to minimi#e the possibility of it

    happening.

    'ntercalated Dis"s

    The most prominent feature of myocardium, and the one that5s absolutelydiagnostic for it, is the intercalated disk. The 'D is a speciali#ed cell2to2cell

    adhesionEcommunications site. 't demarcates the beginning of one myocyte and the

    end of the ne&t, and information is pass across it from cell to cell. These structures

    are found only in cardiac muscle.

    The 'D5s are vital to normal myocyte function and understanding how they are put

    together is important. 'n these special areas there5s a considerable degree of

    interdigitation of myocyte plasma membrane, but there is noactual fusion of

    cytoplasm. The speciali#ed apparatus of the 'D allows myocytes to act as ifthey

    were a true syncytium there is in fact no actual cytoplasmic continuity these are

    distinct and individual cells.

    The true nature of the 'D wasn5t fully understood until the transmission electron

    microscope was available for their study. This instrument revealed that 'D5s are

    physically held together by large numbers of desmosomes, with gap !unctions

    between the myocytes forming a region that permits electrical communication in the

    form of ion flu&es. This flu& maintains the coordination of the waves of contraction

    between one cell and the ne&t.

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    'ntercalated discs are readily seen on slide 9@9, in regions where the myocardium is

    cut longitudinally 0they won5t be visible in cross sections1. Since the 'D5s are located

    at the ends of the cell, and since that5s where the first and last sarcomeres of any

    given myofibril are, you can thin" of the 'D as a sort of thic"ened %terminal

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    &amine the lining of the chambers on slide /. The entire cardiovascular system,

    including all chambers of the heart and all blood vessels, is lined with a simple

    suamous epithelial covering.This lining is among the most metabolically active

    tissues in the body. 'n blood vessels, this is 0by convention1 called %endothelium,%

    but in the heart the special term endocardiumis used instead. 0There isn5t a nic"el5s

    worth of difference between them, but the terminology is so entrenched it couldnever be eradicated.1 'n any

    event, %endocardium% is an apt

    name because it5s literally

    %inside the heart.% The

    endocardium covers the valve

    cusps, too, as you can easily

    verify on this slide. 'f you

    traverse the thic"ness of the

    myocardium and e&amine the

    outersurface of the wall, you

    will see a similar 0though lesswell defined1 layer of simple

    suamous epithelium. Again, by

    convention, this has a special

    name it5s 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. 'tcorresponds to the peritoneum of the

    abdominal cavity in origin 0from the

    mesoderm lining the embryonic coelom1

    and function 0to allow the heart to move

    freely in its cavity without adhesion to

    surrounding structures1. 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 sub2epicardial CT layer, mainly

    reticular fibers.

    ?ur"in!e ibers

    Slide 9F@ is another chun" of myocardium. Most of the features of myocardium are

    visible, but this section also shows ?ur"in!e fibers0;ohannes vangelista von

    ?ur"in!e, /GFG2/F=, a C#ech anatomist, physiologist and microscopist1. These fibers

    are part of the conducting system of the heart.

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    The conducting system transmits the signal to contract from its origin in the

    %pacema"er% to the rest of the myocardium. The ?ur"in!e fibers are not neurons.

    They5re speciali#ed muscle cells, and they loo" li"e it. They can be distinguished

    from the myocardium by their si#e 0they5re much larger than regular myocytes1 and

    by their

    staining0they tend

    to be more

    lightly

    stained

    than true

    myocardium1. 4ith the ?AS stain they5re strongly positive, since they contain large

    amounts of glycogen.

    ?ur"in!e fibers are not contractile, at least not to any significant e&tent. They

    contribute nothing to the force generation of myocardium. The cells of the ?ur"in!e

    fibers are organi#ed into several tracts that lead away from the atrioventricularnode and form nerve2li"e structures. There5s a bundle along both sides of the

    septum, one distributing to the right ventricle and one to the left ventricle. The

    ?ur"in!e fibers are larger and less well stained than the ordinary contractile

    myocardium because of the relatively protein2poor cytoplasm, which doesn5t ta"e up

    the eosin stain very well. Hence the fibers loo" pale and washed out in comparison

    to the protein2pac"ed myocytes. ?ur"in!e fibers do have some of the other features

    of cardiac myocytes, including a centrally2located single nucleus, and even

    intercalated dis"s.

    -ut despite their strictly conductive function, they are muscle cell derivatives, so

    you5re uite li"ely to see striations and other indications of %contractility% if youe&amine them at higher power. There should be two areas of ?ur"in!e fibers in your

    slide, one at each edge of the section. 'n this slide also note the ramifications of the

    muscle, and the intercalated discs.

    The ?ur"in!e fibers ramify into the mass of ventricular myocardium, and from their

    termini the signal is spread through the 'D system. *ecall that all muscle tissue is

    %e&citable% and this property can be used to carry information. That5s what5s

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    happened here. 4hat is a secondary function in contractile myocytes has been

    transmuted into the main role of these not2uite2muscle cells. The fibers run

    through the sub2endothelial connective tissue they can5t generate any force because

    internally they have only a few disorgani#ed actin and myosin fibrils, nothing li"e

    the very regular arrays of sarcomeres in the contractile cells.

    Slide =, shown at left, is

    interesting because it contains the

    atrioventricular node, the

    beginning of the ?ur"in!e fiber

    conduction apparatus.

    This image is ta"en 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 0probably the aorta1 andbelow that a triangular2shaped

    region, the trigona fibrosa. The

    trigona fibrosa is part of the so2

    called %cardiac s"eleton,% the

    dense fibrous CT to which the

    intercellular collagen networ" is

    ultimately anchored. 'f you loo"

    at this structure at high

    magnification it5s very similar in

    appearance to cartilage. The

    trigona fibrosa lies between theatrial 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 'C(. 4hen

    force is e&erted by contraction of

    myocardium, this is the %#ero

    point% against which the force is e&erted. 4hen contraction occurs, the different

    directions of muscle tracts impart a compressive, twisting motion to the heart

    grossly 0you will be able to see this in surgery1. The motion has been compared to

    %wringing out% a wet cloth, and it serves the same function: e&pulsion of the

    ma&imum amount of blood per stro"e. Contraction of the myocardial cells leads to

    reduction in volume of the heart chambers, and conseuently to e!ection of all 0or

    almost all1 of the blood therein. 'mpairment 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 A23 node. 't loo"s li"e a bit of atrial myocardium, splon"ed up

    against the denser and more massive ventricular type. 0This slide also gives you a

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    good comparison of the two types of myocardium side by side1. This isn5t the

    %pacema"er,% 0that5s the sino2atrial or S2A node1 but it receives signals from it and

    transmits them via the conducting fibers to the mass of ventricular myocardium.

    Histologically it loo"s li"e the rest of the atrial myocardium, and in the absence of

    the landmar"s used here, would be very difficult to identify in a section.

    3alves of the Heart

    Slide /6@ is of a valve cusp. >oo" at it at low magnification and at high

    magnification and e&amine 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 wor"ed into it as well. The outer surfaces, e&posed to

    the blood flow, are covered with the epithelium of the endocardial lining, li"e the

    rest of the system. (ote that there5s an e&cursion of myocardium into the base of the

    valve cusp this terminates before the end of the cusp. 't5s continuous with the

    myocardium of the ventricle.

    -lood Supply

    ' hope it hasn5t escaped your notice that the entire mass of the myocardium is shot

    through with blood vessels of varying si#es and shapes, and that there5s an e&tensive

    degree of vasculari#ation. 'n 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 bac" and classify some of these.

    Arteries

    >et us now leave the pumping station and e&amine some of the pipes: the arteries

    and veins. *emember the definition: arteries pump blood away from the heart

    veins carry blood towards the heart. (o other criteria for classification of a vessel as

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    one or the other e&ists.Capillaries are those small vessels within the tissues from

    which o&ygen transfer ta"es 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 bul"

    of the blood to regions of the body where it5s to be distributed. &amples include theaorta, subclavian, and pulmonary arteries. )nce the blood has reached the region of

    distributionsay, the limbsit will be handled by smaller 0but still fairly large1

    distributing or muscular arteries, which send it to sub2regions.As the distribution

    area gets more and more limited the arteries become smaller. 'n very local areas you

    will see small arterioles, essentially mini2arteries with a wall considerably less

    muscular than the larger ones %upstream.%

    lastic or Conducting Arteries

    lastic arteries are constructed li"e fire hoses. *ightly so, because they have the

    same function: to carry a stream of liuid under high pressure. Hence they5redesigned to minimi#e internal friction and flow resistance and to ma&imi#e the

    strength of the wall.

    The lumen is lined with a thin suamous epithelial layer 0the tunica intima1 which

    may be li"ened to the rubber bore of the hose li"e the rubber, it offers a smooth and

    unimpeded passage for the flow of blood. The tunica mediais a region of elastic and

    collagen fibers. lastic arteries 0the aorta most of all1 must withstand an enormous

    head of pressure to pump against the peripheral systemic resistance: conseuently

    the wall is heavily reinforced to prevent bursting, !ust 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 stretchpermitted. 0'n some disease or deficient nutritional statesfor e&ample lathyrism, a

    copper deficiency caused by certain plantsthe wall may be wea"ened, resulting in

    an aneurysmwhich may lea", or burst with fatal effects.

    To complete the analogy, the collagenous tunica adventitiaof the aorta is the fabric

    covering on the outside of the hose. ;ust 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 86 is a fine e&ample of an elastic or conducting artery. 't is, in fact, a section ofthe aorta. The section on this slide is stained with the 3erhoeff5s stainfor elastic

    fibers. 't renders these fibers blac", 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 0see below1. The

    amount of elastic fiber infiltration is so great that no internal or e&ternal elastic

    laminae can be identified.

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    This is a section of the aorta, a nice e&ample of an elastic artery. The elastic fibers

    normally aren5t easily seen in an H preparation li"e 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 mar"ed by arrows. There5s a large clot of blood

    in the field. The bore of this vesselli"e the rest of the cardiovascular systemis

    lined with a simple suamous epithelium.

    The innermost of the numerous elastic layers in this artery has its own name: the

    internal elastic lamina.'t5s the one right up against the tunica intima. The internal

    elastic lamina is much more easily seen in muscular arteries 0see below1 than in the

    elastic ones, though. 'n an elastic artery li"e this one it gets lost in the %bac"ground%

    of do#ens of similar layers.

    As is true of most elastic arteries, the tunica adventitia on this one is a relatively

    small contributor to the wall5s thic"ness.

    There5s more than elastic fibers in the wall of this type of artery. -etween the elasticlayers there is a considerable amount of collagen and some smooth muscle,

    permitting the artery to e&pand under pressure and recoil to original diameter when

    the pressure drops again. Collagenous components in the wall prevent over2

    e&pansion and resist bursting of the vessel.

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    +sing stains specific for elastic fibers reveals how e&tensive the reinforcement of the

    wall can be. Here are two e&amples, again from the aorta. The one at left has been

    stained with a combination of the 3erhoeff and the 3an Iieson stain at about B66&,

    the e&tent of the elastic component 0in blac"1 and the collagenous reinforcementbetween the elastic fibers is easily visible. This field is from the outer edge of the

    tunica media: the almost2completely red area at the bottom is the tunica adventitia.

    At right, the 3erhoeff stain has been used and at about @66& it clearly show elastic

    fibers but not the collagen or smooth muscle, which is unstained by this method.

    3ery large elastic arteries have their own internal blood circulation system and

    nervous supply. They have to: the fibroblasts and other cells that "eep the wall in

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    good shape are so far from the blood supply that diffusion won5t 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 C(S can control blood pressure

    and initiate contractions when needed. The vasa vasorum0and in the case of nerve

    fibers, the nervi vasorum1 i.e., the %vessels of the vessels% and the %nerves of the

    vessels% are a constant feature of big vessels.

    Muscular Arteries

    )nce you get

    the blood out

    to the ma!or

    regions of

    the body,

    there5s a

    transition in

    the structureof the

    arterial wall.

    The

    proportion

    of elastic

    fibers

    decreases,

    and the

    proportion

    of smooth

    muscleincreases.

    Some elastic

    fibers and

    collagen

    fibers will

    always be

    present, but

    eventually

    the great

    bul" of the tunica media will be smooth muscle, and at that point we5re dealing with

    muscularor distributingarteries, whose function is to %distribute% blood supply to

    their regions of responsibility, such as a limb. The artery on slide /68, at right, is a

    dandy e&ample of what a muscular or distributing artery should loo" li"e. This is

    the femoral artery the femoral vein 0see below1 and the femoral nerve are there as

    well. There may be a branch point off this artery on your slide.

    (ote that the wall of the artery is mostly tunica media, and that this tunic is almost

    entirely smooth muscle.lastic CT is present, to be sure, but not nearly to the e&tent

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    that it is in the elastic arteries. There is also a noticeable internal elastic lamina,

    stained bright pin", and a less well defined e&ternal elastic lamina, though at the low

    magnification of this image it5s hard to ma"e 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 !ust

    barely visible in this image. 't5s the undulating pin" line immediately below thelining endothelium. 't mar"s the innermost limit of the muscular tunica media. The

    e&ternal elastic lamina is less regular and can5t be made out at all. (ote the

    prominent collagenous tunica adventitia here. 'n muscular arteries the tunica

    adventitia is often most of the wall5s overall thic"ness. The e&ternal CT investment

    anchors this artery to the surrounding CT.

    The internal elastic lamina is most easily seen in smaller muscular arteries. 'n theseit usually stands out as a bright pin" undulating band !ust below the lining epithelial

    cells and their supporting CT. 'n 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.

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    'n the e&ample at left, the '> is indicated this e&ample also clearly shows the

    separation of the lining epithelium from the '> by the sub2endothelial CT. A few

    layers of smooth muscle constitute the tunica media in this small artery.

    The smooth

    muscle of the wallof distributing

    arteries ma"es

    them very

    e&tensible, and

    also provides for a

    counter force to be

    e&erted against

    the pressure of

    filling. As the

    vessel e&pands the

    smooth musclecells are stretched:

    in reaction to this

    they begin to

    contract. The

    pea" 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. (ervous input can also

    control this to some e&tent, independent of the mechanical force of stretching. As

    distance from the heart increases, the force reuired to dampen the oscillations isless, and smaller arteries can handle it the interval between pea"s also lengthens

    and there is a much more uniform flow rate.

    't5s

    useful to

    put the

    two

    ma!or

    arterial

    types

    side by

    side,

    stained

    to reveal

    the wall

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    components. )n slide 86, this has been done using the 3erhoeff stain. A small

    muscular artery is near the large elastic one, to ma"e the point that as the artery

    si#e 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 bac"ground of the muscular

    tunica media.

    't5s possible to combine several staining methods to show allthe ma!or components

    of the wall, including the smooth muscle. 'n the image below, the 3erhoeff and

    Masson5s stains have been combined. lastic tissue is stained blac" smooth muscle

    is red and collagen is green. The tunica media 0TM1 is almost entirely muscle, with

    a few minor strea"s of green collagen in it. The inner elastic lamina 0'>1 stands out

    prominently and the outer elastic lamina 0)>1 demarcating the end of the tunica

    media is also easily visible. The tunica adventitia 0TA1 is a mi&ture of green collagen

    fibers and blac" elastic fibers interwoven with each other to provide strength and

    resilience.

    Slide F 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 thic" CT tunica adventitia. There are also valves present.

    Arteries 'n >ongitudinal Section

    Since arteries are pretty common in almost any microscope slide you5ll loo" at, it5s

    important to be able to recogni#e them when they aren5t cut in cross section, as all

    the previous e&amples have been.

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    Here are two e&amples from tissue sections. The left section is from slide 8B the

    right from slide /BF. Since the smooth muscle in the wall of an artery is oriented

    with the long a&is of the smooth muscle cells aroundthe long a&is 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 crosssection. Compare these

    profiles 0both at about B66&1 to the sections of smooth muscle in &ercise /6, andyou5ll be able to ma"e out the boundaries of the smooth muscle easily.

    Aneurysms

    Sometimes the wall of an arteryespecially a big one li"e the aorta, which is

    sub!ected to all or nearly all the pressure the heart can generateis wea"ened by

    disease, malnutrition, age, or some other affliction. +nder a good head of pressure,

    http://education.vetmed.vt.edu/Curriculum/VM8054/Labs/Lab12b/Lab12b.htm#%23
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    blood begins to dissect the layers of reinforcing material, separating them and

    causing the side of the artery to bulge, e&actly as a garden hose does when its wall is

    wea"ened. This bulge is an aneurysm. Chronic high blood pressure ma"es this more

    li"ely, though an aneurysm can occur even when systemic blood pressure is lower

    than normal. (eedless to say, an aneurysm that bursts will really spoil your plans

    for the wee"end. 'f it5s a big one that blows out, such as the aorta, death will berapid.

    3eins

    3eins are

    those

    vessels

    leading

    blood bac"

    towards

    the heart.As a rule,

    they have

    much

    thinner

    walls than

    arteries

    do, though

    in cross

    sectional

    area

    they5reusually

    larger

    than the

    corresponding artery, because they have to carry the same volume of blood at a

    lower pressure. Since veins are on the post2capillary side of the circulatory loop,

    operating at much lower pressures, there5s less need for burst resistance. Thin walls

    are also important because much of the pressure that drives blood through veins is

    generated notby the heart, but by contraction of the muscles in the region of the

    vein. This %suishes% the blood bac" through the vein. -ecause they have low

    pressures, some veins have venous valves in them to prevent bac" flow. This is

    especially true of medium si#ed veins in the e&tremities, as they have to lift bloodagainst gravity.

    0'f 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 bac" to the heart. Since the heart can only put out what it ta"es

    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

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    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 "eel over and uite literally %drop out% of the

    ran"s as they

    faint.1

    3enous 3alves

    3eins of a

    certain si#e

    usually have

    valves to

    prevent bac"

    flow. 3eins of

    this type are

    usually found

    in thee&tremities, the

    valves allowing

    blood to move

    bac" towards

    the heart with

    less effort.

    3enous

    pressures are

    so low, and the

    propelling

    force has tofight 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 bac" down when pressure slac"s off.

    The venous valve has a great deal of structural similarity to the valves of the heart.

    There5s a CT core with epithelium on both sides. Two valve flaps meet in the center

    of the vessel. $ou can find such valves on slide F, and when you do, note especially

    the direction of the valve: it5s designed to permit blood to flow in one direction only.

    ?ooling of blood occurs on the superior side of the valve flaps. 'n animals with long

    lives and vertical postures, the continual stress that the weight of this blood imposes

    sometimes causes outpouching of the vein. 'f there is a wea"ening 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 wor" on their feetdevelop in later life.

    This image from slide F shows a fairly large vein, cut more or less in cross section.

    The flaps on both sides of one of its valves are visible.

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    Capillaries

    Capillaries are small vessels, whose walls are thin enough to allow the diffusion of

    nutrients, o&ygen and carbon dio&ide across them.They are %where the action is% in

    gas and nutrient e&change: with a few e&ceptions, no cell of the body is very far

    from one, because access to the blood is an absolute reuirement: cell death wouldresult from ano&ia or the loss of nutrient and waste transport.

    Capillaries are by far the most numerous class of vessels, though they5re 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 %bul"% transfer of materials

    between the blood and the tissue it served isn5t needed, such as in muscles. Closed

    capillaries can move material in and out using a process of seuential endocytosisand e&ocytosis, as well as simple diffusion for small molecules.

    enestrated 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 "idney. They don5t demonstrate the %transcytosis% activity of the

    closed type, because bul" movement of ions, hormones, nutrients, etc. through the

    pores is ample

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    Here5s a scanning

    electron micrograph of

    a capillary running

    alongside muscle

    fibers. This is a closed

    capillary. This stri"ingimage shows the

    appro&imate si#e of a

    capillary very well:

    they5re !ust about the

    same si#e as the

    erythrocytes that flow

    through them, maybe

    /6 7m in luminal

    diameter. 'n this

    picture you can see an

    erythrocyte pee"ingout at the bro"en end,

    and the anchoring

    fibrils of connective

    tissue that attach it to

    the surrounding muscle mass 0*1.

    Compare this to the diagram above: the wall is made of very thin suamous cells

    which are sealed together at the edges by desmosomes in other words, there5s only a

    tunica intima, and not much of that beyond the lining epithelium and the basement

    membrane it rests on. 'n a section it5s normal to see parts of several of the mural

    cells in the same plane, since there5s uite a bit of interdigitation between theirregular borders of ad!acent cells.

    $ou may also see a second cell type, the pericyte. Technically this isn5t part of the

    capillary wall. ?ericytes %embrace% the wall but never ma"e contact with the blood,

    as the mural cells do they5re believed to have some sort of contractile function.

    They5re so closely associated with the mural epithelium they share a basement

    membrane with it.

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    Despite

    their small

    si#e,

    capillaries

    are so

    numerousthat

    they5re

    easy to

    find. They

    often will

    have blood

    cells in

    them,

    which also

    ma"es

    them easyto spot.

    This one is

    from

    s"eletal

    muscle on slide /B. S"eletal 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 !ust large enough to accommodate them

    0double arrow1. A white blood cell 0probably a neutrophil1 is visible in the left end of

    the field. As it happens there are no capillary cell nuclei nor pericyte nuclei visible,

    they5re 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 ma"e out.

    The capillary at right is

    from white fat. 't5s 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. 't shouldn5t

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    therefore be too surprising to find that connective tissues, which are also of

    mesodermal origin, are usually well supplied with blood vessels. That5s the case

    here: white fat is a connective tissue, and the fibrous CT that separates each fat cell

    0the clear spaces in this image1 from the others is a second type of CT. This

    capillary5s 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. 'nyoung animals, which haven5t had time to accumulate the %wear and tear% pigment

    lipofuscin in their fat cells 0see &ercises @ and 8 for a discussion of lipofuscin1 the

    presence of so many capillaries give the fat a pin"ish2white color. 'n albinos, animals

    that lac" melanin, it5s the blood circulating in capillaries of the eye that5s the source

    of its pin" color.

    A fenestrated capillary has %holes%

    in it the word comes fromfenestra,

    the >atin for %window.% These holes

    are actually pores, places where the

    plasma membrane is perforated, toallow for the movement of bul"

    materials from the capillary lumen

    to the e&terior, 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. enestrated

    capillaries are also found in the

    "idney, again a place where uic"transit between two compartments is

    the design criterion.

    't5s not possible to see the

    fenestrations in a light microscope

    preparation. To appreciate the

    nature of these pores, an electron microscopic image is reuired, because they5re

    below the level of the light microscope5s resolution. Such an image is provided here,

    at about F6,666 diameters. This e&ample is from the "idney, but similar capillary

    profiles could be found anywhere fast movement of large molecules is important to

    normal function. The capillaries of the glomerulus 0the tuft of blood vessels that fills

    the renal corpuscle1 use hydrostatic pressure from the arterial supply to filter blood

    plasma through the pores in the formation of urine 0for details, see &ercise @81. As

    this image ma"es 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 -owman5s space. *emember, the imageshown here is

    two2dimensional, but the cellisn5t. The %brea"s% indicated are really more or less

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    circular perforations in the plasma membrane, and they have depth they lie on both

    sides of the plane of the section.

    Sinusoids

    There5s onemore category

    of blood vessels

    to be dealt

    with. These are

    somewhat li"e

    capillaries, and

    might be

    considered as a

    sub2set.

    Sinusoidsare a

    form of large,irregular

    capillary2type

    vessel they

    have the thin,

    intima2only

    wall

    construction of

    a capillary and

    may be

    fenestrated.

    Sinusoids arefound where

    slow flow,

    intimate contact between blood and tissue, and rapid e&change of materials are

    reuired. $ou5ll find them on slide /@8 in the liver, between plates of hepatic cells.

    Sinusoids are

    flattened and

    irregular in shape,

    as the image at

    right ma"es clear.

    This scanning M

    picture is a plastic

    cast of the

    sinusoids in the

    placenta of a goat,

    another place

    where slow flow

    and efficient

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    transfer of materials is important to normal function. 'n 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.