Comparative anatomy of the Circulatory System of

vertebrates
Comparative Anatomy of Vertebrate Hearts
Function: The heart is a hollow muscular organ that rythmically contracts and
relaxes. During each contraction-relaxation cycle, blood is drawn from the veins into a
thin walled collecting chamber, the atrium, and is then passed to a second thick walled
chamber, the ventricle, which forceably contracts to distribute the blood to
the arteries. Backflow is prevented by one-way valves.

Note that the partial septum in the reptile ventricle becomes a complete divider in
birds and mammals.
In the image above, you can see the progressive changes in the heart between
ancestral vertebrates, the fishes, and the most derived forms, the birds and mammals.
Fish have a simple two chambered heart which is, in essense, just a thickening of a
section of the circulatory system, and the blood flows in a single circuit from heart to
gills to body and back to the heart. Starting with the amphibians, the first of the
vertebrates with lungs, the circulatory system adds a second loop or circuit. This
design has the blood flow through the heart twice on each trip around the system, once
on the way to the lungs and once on the way back from the lungs, giving it an extra
boost. This is called double circulation. In amphibians, with two atria but only a
single ventricle, this results in the mixing of deoxygenated and oxygenated blood, but
amphibians also gather oxygen through their moist skin, so this inefficiency is not
critical. Beginning with the reptiles, a septum or wall develops that partly divides the
deoxygenated from the oxygenated blood in the ventricle, and this is important
because reptiles, with a watertight skin, rely entirely on their lungs for oxygen.
Reptiles also have the unique ability to redirect or shunt blood leaving the heart back
through the heart without passing through the body circuit, and to shunt deoxygenated
body blood back through the body without going to the lungs. The purpose of this
shunt (see the purple vessels in the figure below) is not entirely understood. The
former is thought to be a way to prioritize oxygenation of the heart during periods of
high exertion, while the latter is believed to be a way to enhance digestion, because of
increased acidity of deoxygenated blood due to carbon dioxide buildup. Among the
extant reptiles, only the crocodilians have fully extended the septum and have a four-
chambered heart, but there is speculation that dinosaurs may have had this innovation
as well. Birds and mammals have the same four-chambered design, which has
increased efficiency because deoxygenated and oxygenated blood cannot mix.
 Red blood vessels carry oxygen-rich blood. Purple vessels carry mixed blood. Blue
vessels carry deoxygenated blood.

Vertebrate Circulatory Systems:

1- transport gases, nutrients, waste products, hormones, heat, & various other materials

2-consist of heart, arteries, capillaries, & veins:

Arteries carry blood away from the heart have muscular, elastic walls terminate in capillary beds

Capillaries have very thin walls (endothelium only) are the site of exchange between the blood and body
cells

Veins carry blood back to the heart have less muscle in their walls than arteries but the walls are very
elastic begin at the end of capillary beds

Heart is a muscular pump (cardiac muscle) contains a pacemaker to regulate rate but rate can also be
influenced by the Autonomic Nervous System
Cartilaginous fishes

singlecircuit heart with 4 chambers: sinus venosus, atrium, ventricle, & conus arteriosus

the sinus venosus receives blood & is filled by suction when the ventricle contracts & enlarges the
pericardial cavity the atrium is a thinwalled muscular sac; an AV valve regulates flow between atrium &
ventricle the ventricle has thick, muscular walls the conus arteriosus leads into the ventral aorta (and a
series of conal valves in the conus arteriosus prevent the backflow of blood)
Teleosts

heart is similar to that of cartilaginous fishes, except a bulbus arteriosus (a muscular extension of the
ventral aorta) is present rather than a conus arteriosus (a muscular extension of the ventricle)

Lungfish & amphibians

 modifications are correlated with the presence of lungs & enable oxygenated blood returning from the
lungs to be separated from deoxygenated blood returning from elsewhere .

1-Partial or complete partition within atrium (complete in anurans and some urodeles)

2-Partial interventricular septum (lungfish) or ventricular trabeculae (amphibians) to maintain

separation of oxygenated & unoxygenated blood

3- Formation of a spiral valve in the conus arteriosus of many dipnoans and amphibians. The spiral valve
alternately blocks & unblocks the entrances to the left and right pulmonary arches (sending
unoxygenated blood to the skin & lungs).

4-Shortening of ventral aorta, which helps ensure that the oxygenated & unoxygenated blook kept
separate in the heart moves directly into the appropriate vessels
Amniotes:

1 Heart consists of 2 atria & 2 ventricles &, except in adult birds & mammals, a sinus venosus

2 Complete interatrial septum

3 Complete interventricular septum only in crocodilians, birds, & mammals; partial septum in other
amniotes

Arterial channels supply most tissues with oxygenated blood (but carry deoxygenated blood to
respiratory organs). In the basic pattern:

1 the ventral aorta emerges from heart & passes forward beneath the pharynx

2 the dorsal aorta (paired above the pharynx) passes caudally above the digestive tract

3 six pairs of aortic arches connect the ventral aorta with the dorsal aortas

Amphibians:

Urodeles most terrestrial urodeles have 4 pairs of arches;

aquatic urodeles typically have 3 pairs (III, IV, & VI)

 Anurans have 4 arches early in development (larval stage); arch VI develops a pulmonary artery (to
lungs) while arches III, IV, & V supply larval gills.

At metamorphosis: aortic arch 5 is lost the dorsal aorta between arches 3 & 4 is lost, so blood entering
arch 3 (carotid arch) goes to the head a segment (ductus arteriosus) of arch 6 is lost so blood entering
this arch goes to the skin & lungs aortic arch 4 (systemic arch) on each side continue to the dorsal aorta
& distributes blood to the rest of the body Oxygenated blood from the left atrium & deoxygenated
blood from the right are largely kept separate in the ventricle by: Ventricular trabeculae Spiral valve in
the conus arteriosus

Reptiles

3 aortic arches in adults (III, IV, & VI)

Ventral aorta no spiral valve but truncus arteriosus is split into 3 separate passages: 2 aortic trunks & a
pulmonary trunk.
As a result: pulmonary trunk emerges from the right ventricle & connects with 6th aortic arches
(deoxygenated blood from right atrium goes to lungs) one aortic trunk comes out of left ventricle &
carries oxygenated blood to the right 4th aortic arch & to carotid arches the other aortic trunk appears
to come out of right ventricle & leads to left 4th aortic arch. So, does the left 4th arch carry oxygenated
blood?

Turtles, snakes, & lizards the interventricular septum is incomplete where right & left systemic arches
(4th) leave the ventricle & trabeculae in that region of the heart form a ‘pocket’ called the cavum
venosum. Oxygenated blood from the left ventricle is directed into cavum venosum, which leads to the
2 systemic arches. As a result, both the left & right systemic arches receive oxygenated blood

Crocodilians ventricular septum is complete but a narrow channel called the Foramen of Panizza
connects the base of the right & left systemic trunks

Role of the Foramen of Panizza in the crocodilian circulatory system: When a crocodilian is above water
and breathing air, the semilunar valve in the right aorta remains closed because of higher pressure in
the left & right aorta (higher than in the right ventricle). As a result, the right aorta receives blood from
the left aorta (so both aortas carry oxygenated blood) and blood from the right ventricle (low in oxygen)
passes only into the pulmonary artery (and goes to the lungs).

Birds & mammals

no mixing of oxygenated & unoxygenated blood; complete interventricular septum + division of ventral
aorta into 2 trunks: Pulmonary trunk that takes blood to the lungs Aortic trunk that takes blood to the
rest of the body Result of modifications: All blood returning to right side of heart goes to the lungs;
blood returning from lungs to the left side of heart goes to systemic circulation.

Avian Circulatory System

Birds have very efficient cardiovascular systems that permit them to meet the
metabolic demands of flight (and running, swimming, or diving). The cardiovascular
system not only delivers oxygen to body cells (and removes metabolic wastes) but
also plays an important role in maintaining a bird's body temperature.The avian
circulatory system consists of a heart plus vessels that transport:

 nutrients
 oxygen and carbon dioxide
 waste products
 hormones
 heat
Birds, like mammals, have a 4-chambered heart (2 atria & 2 ventricles), with complete
separation of oxygenated and de-oxygenated blood. The right ventricle pumps blood
to the lungs, while the left ventricle pumps blood to the rest of the body. Because the
left ventricle must generate greater pressure to pump blood throughout the body (in
contrast to the right ventricle that pumps blood to the lungs), the walls of the left
ventricle are much thicker & more muscular.

Birds tend to have larger hearts than mammals (relative to body size and mass). The
relatively large hearts of birds may be necessary to meet the high metabolic
demands of flight. Among birds, smaller birds have relatively larger hearts (again
relative to body mass) than larger birds. Hummingbirds have the largest hearts
(relative to body mass) of all birds, probably because hovering takes so much energy

Avian hearts also tend to pump more blood per unit time than mammalian hearts. In
other words, cardiac output (amount of blood pumped per minute) for birds is
typically greater than that for mammals of the same body mass. Cardiac output is
influenced by both heart rate (beats per minute) and stroke volume (blood pumped
with each beat). 'Active' birds increase cardiac output primarily by increasing heart
rate. In a pigeon, for example (Butler et al. 1977):

Rest Active Increase

Heart rate 115 beats/min 670 beats/min 5.8x
Stroke volume 1.7 ml 1.59 ml 0.9x
Cardiac output 195.5 ml/min 1065 ml/min 5.4x
Oxygen consumed 20.3 ml/min 200 ml/min 10x

In general, bird hearts 'beat' at somewhat lower rates than mammals of the same size
but pump more blood per 'beat.' Among birds, heart rate varies with size:

Species Resting heart rate 'Active' heart rate

Turkey 93 -
Herring Gull 130 625
American Robin 570 -
Blue-throated Hummingbird - 1260

Source: Welty & Baptista. 1988. The Life of Birds. Saunders College Publishing, New York.

Blood pumped by the avian heart enters the blood vessels. The main types are:

 arteries - carry blood away from the heart & toward the body cells
 arterioles - 'distribute' blood (that is, direct blood where needed with more
going to active tissues & organs & less to less active tissues & organs) by
vasodilating & vasoconstricting
 capillaries - exchange of nutrients, gases, & waste products between the blood
& the body cells
 venules (small veins) & veins- conduct blood back to the heart

Some of the major arteries in the avian circulatory system:

Carotids deliver blood
to the head (& brain).

Brachials take blood to

the wings.

Pectorals deliver blood

to the flight muscles
(pectoralis).

The systemic arch is

also called the aorta &
delivers blood to all
areas of the body except
the lungs.

The pulmonary arteries

deliver blood to the
lungs.

The celiac (or

coeliac) is the first
major branch of the
descending aorta &
delivers blood to organs
& tissues in the upper
abdominal area.

Renal arteries deliver

blood to the kidneys.

Femorals deliver blood

to the legs &
the caudal artery takes
blood to the tail. Source: http://numbat.murdoch.edu.au/Anatomy/avian/avian3.html

The posterior
mesenteric delivers
blood to many organs
& tissues in the lower
abdominal area.
Some major veins in the avian circulatory system:

The jugular
anastomosis allows
blood to flow from right
to left side when the
birds head is turned &
one of the jugulars
constricted.

The jugular veins drain

the head and neck.

The brachial veins drain

the wings.

The pectoral veins drain

the pectoral muscles and
anterior thorax.

The superior vena

cavae (or precavae)
drain the anterior regions
of the body.

The inferior vena

cava (or postcava) drains
the posterior portion of
the body.

The hepatic vein drains

the liver.

The hepatic portal

Source: http://numbat.murdoch.edu.au/Anatomy/avian/avian3.html vein drains the digestive
system.

The coccygeomesenteric
vein drains the posterior
digestive system &
empties in the hepatic
portal vein.

The femoral veins drain

 the legs.

The sciatic veins drain

the hip or thigh regions.

The renal & renal

portal veins drain the
kidneys.

The heart pumps & the vessels carry, of course, blood. Avian blood:

 consists of plasma + formed elements

o plasma is largely water (~85%) plus lots of protein (~9-11%); other
constituents of blood include glucose (blood glucose levels in birds are
greater than in mammals; about 200-400
mg/dl), amino acids, waste products,
hormones,antibodies, & electrolytes.
o the formed elements include red blood
cells (or erythrocytes), white blood cells
(or leucocytes), and thrombocytes
 bird red blood cells (shown to the
right), unlike those of mammals,
are elliptical in shape and nucleated. In most species, red blood
cells are about 6 x 12 microns in size (mammalian RBC's are
typically 5.5 - 7.5 microns in diameter). Typical concentrations
are 2.5 to 4 million/cubic mm. Avian red blood cells have a
lifespan of 28-45 days (shorter than mammals, e.g., about 120
days in humans). Red blood cells contain hemoglobin, the
molecule responsible for transporting oxygen throughout the
body, and are produced in the bone marrow. However, many bird
bones are pneumatic (penetrated by air sacs) and do not contain
marrow. Hemopoietic bone marrow (red-blood-cell-producing
marrow) is located in the radius, ulna, femur, tibiotarsus, scapula,
furcula (clavicles), pubis,and caudal vertebrae.

 Differences in the red blood cells of birds and mammals -- Mammals, which
had developed an aerobic metabolism, emerged in the Triassic, when the
oxygen content in the atmosphere was by approximately 50% lower than
current levels and even lower than in the Jurassic period (when birds evolved).
Under these conditions, natural selection favored the loss of nuclei in the red
blood cells of mammals (making the cells smaller and allowing capilaries to
become even smaller in diameter) and change to a biconcave shape (increasing
 the amount of surface area and enhancing diffusion into and out of the red
blood cells). Birds, with their efficient respiratory system, evolved during the
Jurassic when the oxygen content in the Earth atmosphere approached the
present level, so there was no selective pressure to eliminate nuclei from their
red blood cells or change in shape (Gavrilov 2013).

 bird thrombocytes (shown above with two red blood cells), also
nucleated, are comparable to the non-nucleated platelets of
mammalian blood. Thrombocytes are important in hemostasis
(blood clotting).
 White blood cells play an important role in protecting birds from
infectious agents such as viruses and bacteria. Birds have several
types of white blood cells:

Avian White Blood Cells

The heterophil is the Monocytes are motile cells Eosinophils make up

The lymphocyte is the most
second most numerous that can migrate using about 2 to 3 % of the WBC
numerous white blood cell.
WBC in most birds. ameboid movements. population of healthy birds.
Lymphocytes are either T-
Heterophils are Monocytes are also The function is these cells
lymphocytes (formed in the
phagocytic and use their phagocytic. is unclear.
thymus) or B-lymphocytes
enzyme-containing
(formed in the bursa of
granules to lyse ingested
Fabricius). B-lymphocytes
materials. Heterophils
produce antibodies; T-
are motile and can leave
lymphocytes attack infected
blood vessels to engulf
or abnormal cells.
foreign materials.

 Simplified diagram of the human Circulatory system in anterior view.

Main Features of the Human Circulatory System

 A liquid, blood, to transport nutrients, wastes, oxygen and carbon dioxide, and
hormones.
 Two pumps (in a single heart): one to pump deoxygenated blood to the lungs and
the other to pump oxygenated blood to all the other organs and tissues of the body
 A system of blood vessels to distribute blood throughout the body
 Specialized organs for exchange of materials between the blood and the external
environment; for example organs like the lungs and intestine that add materials to the
blood and organs like the lungs and kidneys that remove materials from the blood and
deposit them back in the external environment

The Heart and Pulmonary System

The heart is located roughly in the center of the chest cavity. It is covered by a protective
membrane, the pericardium.

 Deoxygenated blood from the body enters the right atrium.

 It flows through the tricuspid valve into the right ventricle. The term tricuspid refers
to the three flaps of tissue that make up the valve.
 Contraction of the ventricle then closes the tricuspid valve and forces open the
pulmonary valve.
 Blood flows into the pulmonary artery.
 This branches immediately, carrying blood to the right and left lungs.
 Here the blood gives up carbon dioxide and takes on a fresh supply of oxygen.
 The capillary beds of the lungs are drained by venules that are the tributaries of
the pulmonary veins.
 Four pulmonary veins, two draining each lung, carry oxygenated blood to the left
atrium of the heart.

Figure 15.3.1.1 Human heart

The above figure shows the human heart, with a schematic view of the pathway of blood
through the lungs and internal organs. Oxygenated blood is shown in red; deoxygenated blood
in blue. Note that the blood draining the stomach, spleen, and intestines passes through the
liver before it is returned to the heart. Here surplus or harmful materials picked up from those
organs can be removed before the blood returns to the general circulation.

The Coronary System

From the left atrium,

 Blood flows through the mitral valve (also known as the bicuspid valve) into the left
ventricle.
 Contraction of the ventricle closes the mitral valve and opens the aortic valve at the
entrance to the aorta.
  The first branches from the aorta occur just beyond the aortic valve still within the
heart.
 Two openings lead to the right and left coronary arteries, which supply blood to the
heart itself. Although the coronary arteries arise within the heart, they pass directly out
to the surface of the heart and extend down across it. They supply blood to the network
of capillaries that penetrate every portion of the heart.
 The capillaries drain into two coronary veins that empty into the right atrium.

DISEASES OF THE CORONARY SYSTEM: ARTERIOSCLEROSIS AND ATHEROSCLEROSIS

The coronary arteries arise at the point of maximum blood pressure in the circulatory system.
Over the course of time, the arterial walls are apt to lose elasticity, which limits the amount of
blood that can surge through them and hence limits the supply of oxygen to the heart. This
condition is known as arteriosclerosis.

Alternatively, fatty deposits, called plaque, may accumulate on the interior surface of the
coronary arteries; this condition is known as atherosclerosis. This is particularly common in
people who have high levels of cholesterol in their blood. Plaque deposits reduce the bore of
the coronary arteries and thus the amount of blood they can carry. Atherosclerosis (usually
along with arteriosclerosis) may limit the blood supply to the heart that during times of stress
the heart muscle is so deprived of oxygen that the pain of angina is created. It triggers the
formation of a clot causing a coronary thrombosis. This stops the flow of blood through the
vessel and the capillary network it supplies causing a heart attack. The portion of the heart
muscle deprived of oxygen dies quickly of oxygen starvation. If the area is not too large, the
undamaged part of the heart can, in time, compensate for the damage.

Coronary bypass surgery uses segments of leg veins to bypass the clogged portions of the
coronary arteries.

The Systemic Circulation

The remainder of the system is known as the systemic circulation. The graphic shows the major
arteries (in bright red) and veins (dark red) of the system. Blood from the aorta passes into a
branching system of arteries that lead to all parts of the body. It then flows into a system of
capillaries where its exchange functions take place.
 Figure 15.3.1.2 Human circulation system

Blood from the capillaries flows into venules which are drained by veins.

 Veins draining the upper portion of the body lead to the superior vena cava.
 Veins draining the lower part of the body lead to the inferior vena cava.
 Both empty into the right atrium.

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