Circulatory

System
BUSTILLO, JOHN GELMARCK B.
VILLANUEVA, CZARAE JANA D.
 Blood

▪ Cells produced by hemopoietic tissues usually enter the circulation to become the peripheral or circulating blood

▪ The plasma is the fluid component and can be thought of as the ground substance of blood, a special connective
tissue.

▪ The formed elements are the cellular components of blood.

▪ Hemoglobin, the major oxygen transport molecule, is excreted by the kidneys if it is left free in the plasma. Thus,
red blood cells function as containers for hemoglobin, preventing its elimination.

▪ White blood cells, or leucocytes, are a second major cellular constituent of the formed elements
▪ . Leucocytes defend the body from infection and disease.

▪ The platelets are a third formed element in the blood. They release factors that produce a cascade of chemical
events leading to the formation of a clot, or thrombus, at sites of tissue damage.
 Arteries, Veins, and Capillaries

Although they vary in size, there are three principal types of blood vessels: arteries, veins,
and capillaries.
▪ Arteries carry blood away from the heart, veins carry blood toward the heart, and
capillaries are the tiny vessels that lie between them.
 Arteries and veins have tubular walls organized into three layers
that enclose a central lumen (figure 12.1)

▪ The innermost layer, the tunica intima, includes the lining of endothelial cells that face the lumen. On the outside is
the tunica adventitia, composed mostly of fibrous connective tissue. Between these two layers is the tunica media,
which differs the most in arteries and veins
▪ Very small arteries and veins are called arterioles and venules.
▪ Smooth muscles form sheets that surround the walls of arteries or veins. Smooth
muscle cells respond to nervous and hormonal stimuli. When they contract, the
caliber of a vessel narrows, a response termed vasoconstriction.
▪ When circular muscle contraction ceases, resident blood pressure forces the vessel to
open and restore or enlarge the size of the lumen, a response called vasodilation.
*Obliquely oriented smooth muscle may also assist vasodilation
Arteries

▪ The structure of arteries varies with their size. ▪ With their elastic walls, these large arteries expand to
receive the sudden injection of blood, which can be felt in
▪ Large arteries have considerable amounts of elastic fibers your wrist or neck arteries as a “pulse.”
in their walls; small arteries have almost none
▪ ” Between contractions, the stretched arterial walls
▪ Large arteries have considerable amounts of elastic fibers elastically recoil, driving this volume of blood smoothly
in their walls; small arteries have almost none along through the smaller arteries and into the arterioles

▪ Arteries function primarily as a supply system that carries ▪ Arterioles direct this blood to local tissues. Humans are
blood away from the heart and out to body tissues. one of the few species prone to arterial disease,
characterized by hardening of the arterial walls and loss of
▪ They also absorb and distribute the sudden surge of blood elastic recoil. As a consequence, afflicted arteries do not
through them when the heart contracts. expand to blunt the sudden pulse of blood, nor do they
move the column of blood along between heartbeats.
▪ Rhythmic contractions of the heart send spurts of blood
into the large arteries. ▪ The heart must work harder, and the smaller arteries and
arterioles experience higher surges of blood pressures.
Not designed for such pressures, these small vessels may
rupture. If this occurs in a critical organ such as the brain,
death may follow.
 Hemodynamics of Circulation

▪ The pressures and flow patterns of the blood ▪ The diastolic pressure constitutes the lowest pressure
circulating through vessels constitute the within the blood vessels, which is reached between
hemodynamics of circulation. heartbeats.

▪ When the ventricles of the heart contract, the peak

force produced is the systolic pressure.
 Veins

▪ Because veins return blood to the heart, they are

collecting tubes. At any one moment, up to 70% of the
circulating blood within the body may reside in veins.

• Veins are also designed to address low blood pressures.

One-way valves that prevent retrograde blood flow are
common within their walls. If veins pass between active
muscles or through parts of the body subjected to
pressure changes (e.g., the pleural cavities containing
the lungs), external forces impinge upon and squeeze
their walls. These supplemental forces contribute to
venous flow, and because of the one-way valves, blood
moves only one way back to the heart (figure 12.3).
Understandably, in veins that pass through body organs
and tissues that offer no induced forces, such as those
within the bones or the brain, one-way valves are absent,
and return of the blood to the heart depends on any
remaining intrinsic pressure and gravity.
 Microcirculation

▪ The specific component of the ▪ Blood can be diverted through shunts that
cardiovascular system that regulates and bypass some regions entirely (figure
supports cell metabolism intimately is the 12.4).
microcirculation.
▪ The precapillary sphincters are little rings
of smooth muscle restricting the entrance
to the capillary beds.
▪ As an animal lowers its head to drink from a stream, blood ▪ Following severe injury or trauma, the microcirculation may fail
pressure within its tissues changes quickly (figure 12.5). to regulate blood distribution. When this happens, a condition
called shock, properly hypotensive shock, results from a
▪ Humans who exercise might complain of “side cramps.” This cascade of events.
results from ischemia, a localized lack of sufficient blood to the
stomach to meet metabolic expectations.
 Single and Double Circulation
▪ Blood travels in one of two general patterns. Most fishes ▪ Amniotes have a double circulation pattern in which blood
have a single circulation pattern in which blood passes only passes through the heart twice during each circuit, traveling
once through the heart during each complete circuit. With from the heart to the lungs, back to the heart, out to the
this design, blood moves from the heart to the gills to the systemic tissues, and back to the heart a second time (figure
systemic tissues and back to the heart (figure 12.6a). 12.6b
 Embryonic Development of the Cardiovascular
System
Most blood vessels arise within embryonic mesoderm (or ▪ Embryonic blood islands yield both blood vessels and
from mesenchyme) almost as soon as this germ layer blood cells, so they are involved in angiogenesis (blood
becomes established vessel formation) and hemopoiesis (blood cell
formation).
▪ Small clusters of mesodermal cells called blood islands
mark the embryonic debut of the cardiovascular system ▪ The sinus venosus is the first chamber to receive
(figure 12.7a–d) returning blood. Blood flows next into the atrium, then
into the ventricle, and finally into the fourth chamber, the
bulbus cordis.
▪ Development of the heart begins when cells leave the splanchnic mesoderm to form a
medial pair of endocardial tubes (figure 12.8a,b).
▪ Cells remaining in the splanchnic mesoderm proliferate, producing a thickened lateral
region, the paired epimyocardium.
▪ Specifically, the fused endocardial tubes form the endothelial lining of the heart, called
the endocardium, and the epimyocardium gives rise to the extensive cardiac muscle of
the heart wall, the myocardium, together with the thin visceral peritoneum covering the
heart’s surface. With these fusions, the basic four chambered embryonic heart is
established (figure 12.8c).
▪ Subsequent folding of the tubular heart twists the heart into different
configurations, but the internal sequence of blood flow remains the same
(figure 12.9).
 Phylogeny of the Cardiovascular System

▪ The vessels of the cardiovascular system are as varied as the diverse organs they
supply. However, these variations are based on modifications of a fundamental plan of
organization common to vertebrates. Because it is usually highly modified in advanced
forms, this fundamental organization of the cardiovascular system is most evident in
primitive vertebrates
▪ Blood leaving the heart first enters an unpaired ventral aorta
and courses forward below the pharynx.

▪ Anteriorly, the ventral aorta divides into the external carotids,

which carry blood into the ventral region of the head.

▪ Before producing these external carotids, however, the

ventral aorta gives off a series of aortic arches, which pass
dorsally within the branchial arches between pharyngeal
slits.

▪ Above the pharynx, these aortic arches meet a paired dorsal

aorta.

▪ Sprouting from the anterior end of the dorsal aorta are the
internal carotids, which carry blood forward into the head
and usually penetrate the braincase to supply the brain.

▪ The dorsal aorta itself, however, carries blood posteriorly

(figures 12.10 and 12.11).
▪ At about the level of the liver, the paired vessels of the dorsal aorta unite to form the unpaired aorta, which distributes
blood to the posterior part of the body and eventually extends into the tail as the caudal artery.

▪ Along the way, the dorsal aorta gives off numerous small parietal arteries to the local body wall, as well as several
major arteries, usually paired, to somatic tissues.
• Paired subclavian arteries supply the anterior appendages (fins or limbs) and usually branch from the dorsal aorta, as
do the caudal iliac arteries, which supply the posterior appendages.
• The gonads receive blood from paired) genital arteries (ovarian or spermatic).
• Paired renal arteries to the kidneys are large, major branches from the dorsal aorta. This ensures that the kidneys
receive blood early in the arterial circuit while blood pressure is still relatively high, a feature of the hemodynamics that
aids renal filtration. Typically in vertebrates, three unpaired arteries depart from the dorsal aorta to supply the viscera.
These are the celiac, supplying the liver, spleen, stomach, and part of the intestines; the anterior mesenteric, supplying
most of the small intestine; and the posterior mesenteric, supplying the large intestine.
▪ A portal system is a vascular pathway that begins in one set of
capillaries and runs to another without coursing through the
heart in between. ▪ Aortic Arches The number of primitive aortic arches and
branchial arches through which they run are still debated. Some
▪ The hepatic portal system begins in capillaries within the wall of ostracoderms had as many as ten pairs of branchial arches and
the digestive tract and runs as the hepatic portal vein to the liver, presumably ten pairs of aortic arches. The number of pairs of
where it empties into the capillaries and blood sinuses of the aortic arches varies in living forms. Lampreys have 8 (figure
liver. 12.12), hagfishes 15

▪ The renal portal system transports blood returning from capillary ▪ The section of the aortic arch delivering blood to the gills is the
beds within the tail or hindlimbs through the paired renal portal afferent artery, and the dorsal section carrying it away is the
veins, which empty into capillaries within the kidneys efferent artery.

▪ In some fishes and certainly in tetrapods, the cardinal veins ▪ The capillary beds between them partially or completely encircle
become less involved in blood return. Instead, the composite, the gills and empty first into the collecting loop, which joins the
prominent postcava (posterior vena cava) arose to drain the efferent artery.
posterior part of the body and the) precava (anterior vena cava)
developed to drain the anterior part of the body. Beginning first
with arterial vessels, let us now follow the major phylogenetic
modifications within the cardiovascular system in detail.
• In chondrichthyans, the first pharyngeal slit becomes reduced but not
lost, forming the small spiracle.
• This vessel constitutes the afferent spiracular artery.
• The dorsal section of the first arch forms the efferent spiracular artery,
which drains this small capillary bed.
• The anterior and posterior halves of each collecting loop are its pretrematic and posttrematic branches,
respectively (figure 12.15a).
• In all lungfishes, the efferent vessel of the most
posterior aortic arch (VI) gives rise to the
pulmonary artery but maintains its connection to
the dorsal aorta via the short ductus arteriosus.
• The short section of dorsal aorta between aortic
arches III and IV, termed the carotid duct, usually
closes at metamorphosis.
• The section of ventral aorta between arches III
and IV becomes the common carotid artery, which
feeds the external carotid (from the anterior
ventral aorta) and the internal carotid (the anterior
section of the dorsal aorta together with the third
aortic arch)
• The carotid body is a small cluster of sensory cells
associated with capillaries, usually located near
the point at which the common carotids branch.
• The last arch (VI) loses its connection to the dorsal aorta because
the ductus arteriosus closes and becomes the pulmocutaneous
artery.
• The other branch is the cutaneous artery, which delivers blood to
the skin along the dorsal and lateral body wall.
• The pulmonary trunk incorporates the bases of the paired sixth
arch and their branches as part of the pulmonary arch to the
lungs.
• The base of the left aortic arch, the left aortic arch (IV) itself, and
the curved section of the left dorsal aorta into which it continues
constitute the left systemic arch.
• The right systemic arch includes the same components on the
right side of the body: the base of the right aortic arch, the right
aortic arch itself, and the arched section of the right dorsal aorta.
 Venous Vessels
The major veins that return blood to the heart are complicated and highly variable. Within each vertebrate group,
the veins compose a few main functional systems that arise embryologically from what seems to be a common
developmental pattern. Before examining the anatomy of veins in each group, we turn first to these basic
systems of venous circulation. In vertebrates with an established double circulation, there are two general
functional systems of venous circulation: the systemic system draining the general body tissues, and the
pulmonary system draining the lungs. Within the systemic system, hepatic portal veins serve the liver, renal
portal veins serve the kidneys, and general body veins drain the remaining systemic tissues.

Systemic System Early in development, three major sets of paired veins are present: the vitelline veins from the
yolk sac, the cardinal veins from the body of the embryo itself, and the lateral abdominal veins from the pelvic
region.
Pulmonary System Many fishes have supplementary airbreathing organs, but only fishes
with lungs possess a pulmonary system. Among living fishes, only dipnoans have true
lungs. If the ancient placoderms had lungs, a possibility mentioned earlier, then the
pulmonary system would have evolved early in vertebrate evolution.

Pulmonary Veins The pulmonary veins return blood from the paired lungs to the heart.
Before entering the heart, they usually unite into a single vein. Embryologically, the
pulmonary vein does not arise by conversion of existing vascular channels. Instead,
numerous small vessels originate separately within and drain the embryonic lung buds.
They then converge into several common vessels that become the pulmonary veins
entering the left atrium.

Fishes The head is drained by the paired anterior cardinal veins and small inferior jugular
veins, which join the common cardinal veins just before they empty into the sinuvenosus of
the heart. The subclavian and iliac veins drain the appendages via the lateral abdominal
vein. Both also join the common cardinal. In most fishes, modification of the posterior
cardinal diverts all returning blood from the tail so that it flows through the kidneys before
emptying into the remaining sections of the posterior cardinal. The hepatic portal vein
transports blood from the digestive tract to the capillaries in the liver. From the liver, blood
flows to the heart via the short hepatic veins (figure 12.23a,b).
• Amphibians In larval salamanders, such as
Necturus, the internal jugular (derived from the
anterior cardinal) and external jugular veins,
together with the small lingual vein from the
tongue, return blood from the head. The large
postcava offers one route for blood returning from
the kidneys.
• The ventral abdominal vein transports blood
primarily from the hindlimbs to the liver sinusoids.
Blood from the tail is offered several alternative
return routes: through the ventral abdominal, the
posterior cardinal, or the postcava via the kidneys.

• Reptiles The internal jugular (derived from the

anterior cardinal), external jugulars, and subclavian
from the forearm are tributaries of the paired
common cardinals. The enlarged and modified
common cardinals are customarily called the
precava in reptiles. The posterior cardinal is
considerably reduced to small azygos veins draining
the inner wall of the thorax
• Birds Short external jugulars join long internal
jugulars (anterior cardinals) to return blood to the
common cardinals, which are modified into the
paired precava. The femoral, caudal, and renal veins
are tributaries of the extensive postcava, which also
receives hepatic veins before entering the heart.

• Mammals Renal portal and abdominal veins are

absent in mammalian venous circulation, but a
hepatic portal vein is present. The cardinal vessels
are substantially modified to produce two major
vessels: the single precava (the superior vena cava in
humans) and the single postcava (inferior vena cava
in humans). These vessels collect blood from the
anterior and posterior parts of the body, respectively,
and return it to the right atrium of the heart.
Hearts
The heart is a pump that moves blood through vessels both by pushing blood
through the circulatory system and also by aspiration—creation of negative pressure that
sucks blood into the heart. In a slow-swimming dogfish shark, the heart can move 7.5
liters of blood per hour; in a resting hen, 24 liters per hour; in a human, 280 liters (about
75 gallons) per hour. In a giraffe, almost 1,200 liters of blood can circulate throughout
the body per hour. If heart rate increases, a response known as tachycardia, these values
can increase fivefold. If heart rate decreases, bradycardia results and these values can
fall precipitously

Basic Vertebrate Heart

Phylogenetically, the heart probably began as a contractile vessel, much like those
found within the circulatory system of amphioxus. In most fishes, the heart is part of a
single circulation. Vessels serving gas exchange in the gills and systemic capillary beds
are in series with each other. The embryonic fish heart consists of four chambers, which
are also in series, so that blood flows in sequence from the sinus venosus, to the atrium,
to the ventricle, and finally to the fourth and most anterior heart chamber, the bulbus
cordis, before entering the ventral aorta. Differences in structure, doubts about
homology, and loose use of terms have led to confusion about nomenclature for this
fourth chamber. We use the term bulbus cordis for this chamber in embryos. In adults,
the ventricle empties into the ventral aorta, or to this intervening fourth chamber, the
bulbis cordis, termed in the adult the conus arteriosus if its contractile walls possess
cardiac muscle, bulbus arteriosus if its elastic walls lack cardiac muscle. Internally, each
may contain various numbers of conal valves.
• Occasionally, the hagfish heart is called a
branchial heart to distinguish it from unique
accessory blood pumps elsewhere in its
circulation (figure 12.25b). These
supplementary circulatory pumps are
sometimes called accessory “hearts,” in
quotation marks because they contract but
usually lack the cardiac muscle of true
branchial hearts (figure 12.25c,d).

• The cardinal hearts lying within the anterior

cardinal veins are like sacs whose pumping
action is initiated by skeletal muscles around
their outer walls. The paired caudal hearts, which
are located in the tail, represent a unique
bloodpumping mechanism among vertebrates.
They are composed of a central cartilaginous rod,
skeletal muscles on the side, and veins between.
Alternative contraction of these muscles bends
the rod back and forth, which presses on the
walls of the vessels and pumps blood into the
caudal vein (figure 12.25d).
• The portal heart is a single, expanded vascular sac that receives venous blood from one anterior and one posterior
cardinal vein, and then it contracts to drive the blood through the liver (figure 12.25c). Only hagfishes have such an
accessory heart in the course of the hepatic portal vein, which elevates blood pressure prior to the blood’s entry into
the liver sinusoids. Furthermore, the portal heart is the only accessory heart to have walls of cardiac muscle like the
cardiac muscle of true branchial hearts.
• A single pair of bulbar valves at the juncture of
the bulbus arteriosus and the ventricle prevents
retrograde.
• The result is a depulsation, or dampening of the
large oscillations in blood flow and pressure
introduced by ventricular contractions.
• Lungfishes The lungfish heart is modified from that of other bony fishes. The first
chamber to receive returning blood is still the sinus venosus. In all three lungfish
genera, the single atrium is partially divided internally by an interatrial septum
(pulmonalis fold) that defines a larger right and smaller left atrial chamber (figure
12.29a).
• In place of the atrioventricular valves is the atrioventricular plug, a raised cushion
in the wall of the ventricle.
• The ventricle is also divided internally, but only partially, by an interventricular
septum.
• Amphibians rely on cutaneous gas exchange (plethodontid
salamanders lack lungs entirely), on gills (many larval forms), on lungs
(most toads and frogs), or on all three modes (most amphibians).
Because the sources of oxygenated and deoxygenated blood vary,
heart structure varies as well. Generally, in amphibians with functional
lungs, the heart includes a sinus venosus, right and left atria divided
by an anatomically complete interatrial septum, a ventricle lacking
any internal subdivision, and a conus arteriosus with a spiral valve
(figure 12.30a).
• The cardiovascular system is perhaps best studied in frogs. The conus
arteriosus of the frog heart arises from a single trabeculate ventricle
(figure 12.30b).
• In lungless salamanders or those with reduced lung function, the
interatrial septum and spiral valve may be much reduced or absent
entirely. Unlike frogs, in which the pulmocutaneous artery branches
give rise to the cutaneous artery, salamanders lack a cutaneous
artery.
• The trabeculae apparently are the structures in the frog ventricle that
separate pulmonary and systemic venous streams of blood flowing
through the heart (figure 12.30b)
• The pulmonary artery and the systemic arches in salamanders arise
from the truncus arteriosus (figure 12.31a,b).

• The two different streams of blood returning from the systemic and
pulmonary circuits of amphibians are kept mostly separate as they pass
through the heart (figure 12.31c).
Reptiles have entered more fully terrestrial environments and
adopted more active lifestyles than have amphibians. The
cardiovascular system of reptiles supports accompanying higher
metabolic rates and elevated levels of oxygen and carbon dioxide
transport.
• It is capable of generating elevated blood pressures, higher
cardiac output, and efficient separation of oxygenated and
deoxygenated bloodstreams.

Chelonian/Squamate Hearts
• In these reptiles, the sinus venosus is reduced in comparison to
amphibians but it retains the same functions.
• The conus arteriosus (or bulbus cordis) appears during early
embryonic development but becomes divided in the adult to form
the bases (trunks) of three large arteries leaving the ventricle: the
pulmonary trunk and the right and left systemic trunks.
• Crocodilian Hearts In many respects, the hearts of alligators and crocodiles are
structurally similar to those of other reptiles.
• One-way lunar valves at the bases of each trunk permit blood to enter the conus but halt
reverse backflow into the ventricle
• A narrow channel called the foramen of Panizza connects the left and right aortic arches
shortly after they depart from the ventricle.
Fetal Circulation in Placental Mammals

• A single umbilical vein carrying oxygenated blood away from the placenta flows to the
liver, where approximately half the blood enters the sinusoids of the liver and the
other half bypasses the liver via the ductus venosus and enters the hepatic vein.
• About 90% of the blood that reaches the pulmonary artery bypasses the lung via the
ductus arteriosus and is diverted instead to the dorsal aorta.
• The foramen ovale, an opening between right and left atria, allows most blood
entering the right atrium to flow directly to the left atrium without first passing
through the lungs.
 Lymphatic System
The lymphatic system is partner with the circulatory • Hydrostatic pressure represents the remaining
system. It aids fluid return to the circulatory system force generated initially by ventricular contraction.
and is engaged in several special functions. • Osmotic pressure results from unequal
Structurally, there are two components of the concentrations of proteins within the arteriole and
lymphatic system: lymphatic vessels and lymphatic outside in the surrounding tissue fluid, so fluid
tissue. moves from the surrounding tissue into the blood.
• Fluid seeps from the blood to bathe the
Lymphatic Vessels surrounding cells. This fluid that has escaped from
Collectively, the lymphatic vessels constitute a the blood capillaries is called tissue fluid.
blind-ended, tubular system that recirculates fluid • The remaining 10%, if not recovered, would build
from the tissues back to the cardiovascular system. up in connective tissues, causing them to swell
The walls of lymphatic vessels are similar to those of with excess fluid, a condition termed edema.
veins and, like veins, lymphatic vessels contain one- • The fluid carried by the lymphatic vessels is lymph.
way valves. • In some vertebrates, such as teleost fishes and
amphibians, lymph “hearts” occur along the route
of return.
Lymphatic Tissue

• A lymph node (figure 12.47b) is a collection of lymphatic

tissue wrapped in a capsule of fibrous connective tissue.
• In reptiles, dilation or expansion of lymphatic vessels, termed
lymphatic cisterns or lymphatic sacs, occurs at locations
usually occupied by true lymph nodes in birds and mammals.
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