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Heart

This article is about the internal organ. For other uses, of deaths.[9][10] Of these more than three quarters folsee Heart (disambiguation).
low coronary artery disease and stroke.[9] Risk factors include: smoking, being overweight, little exercise, high
pressure, and poorly controlled
The heart is a muscular organ in humans and other an- cholesterol, high blood [11]
diabetes, among others.
Diagnosis of CVD is often
imals, which pumps blood through the blood vessels of
done
by
listening
to
the
heart-sounds
with a stethoscope,
[1]
the circulatory system. Blood provides the body with
ECG or by ultrasound.[3] Specialists who focus on disoxygen and nutrients, and also assists in the removal of
metabolic wastes.[2] The heart is located in the middle eases of the heart are called cardiologists, although many
specialties of medicine may be involved in treatment.[10]
compartment of the mediastinum in the chest.[3]
In humans, other mammals and birds the heart is divided
into four chambers: upper left and right atria; and lower
left and right ventricles.[4][5] Commonly the right atrium
and ventricle are referred together as the right heart and
their left counterparts as the left heart.[6] Fish in contrast
have two chambers, an atrium and a ventricle, while reptiles have three chambers.[5] In a healthy heart blood ows
one way through the heart due to heart valves, which prevent backow.[3] The heart is enclosed in a protective sac,
the pericardium, which also contains a small amount of
uid. The wall of the heart is made up of three layers:
epicardium, myocardium, and endocardium.[7]

1 Structure

The heart pumps blood through both circulatory systems.


Blood low in oxygen from the systemic circulation enters the right atrium from the superior and inferior vena
cavae and passes to the right ventricle. From here it is
pumped into the pulmonary circulation, through the lungs
where it receives oxygen and gives o carbon dioxide.
Oxygenated blood then returns to the left atrium, passes
through the left ventricle and is pumped out through the
aorta to the systemic circulationwhere the oxygen is
used and metabolized to carbon dioxide.[2] In addition the
blood carries nutrients from the liver and gastrointestinal
tract to various organs of the body, while transporting
waste to the liver and kidneys. In the healthy organism each heartbeat causes the right ventricle to pump
the same amount of blood into the respiratory organ as
the left ventricle pumps to the body. Veins transport
blood to the heart and carry deoxygenated blood - except
for the pulmonary and portal veins. Arteries transport
blood away from the heart, and apart from the pulmonary
artery hold oxygenated blood. Their increased distance
from the heart cause veins to have lower pressures than
arteries.[2][3] The heart contracts at a resting rate close to
72 beats per minute.[2] Exercise temporarily increases the
rate, but lowers resting heart rate in the long term, and is
good for heart health.[8]

Real-time MRI of the human heart

The heart is situated in the middle of the mediastinum behind the breastbone in the chest, at the level of thoracic
vertebrae T5-T8. The largest part of the heart is usually
slightly oset to the left side of the chest (though occasionally it may be oset to the right) and is felt to be on
the left because the left heart is stronger, since it pumps
to all body parts. Because the heart is between the lungs,
the left lung in turn is smaller than the right lung because
it has to accommodate the heart.

The heart is supplied by the coronary circulation and is


enclosed in a double-membraned sacthe pericardium.
This attaches to the mediastinum, providing anchorage
[13]
The back surface of the heart lies near
Cardiovascular diseases (CVD) are the most common for the heart.
cause of death globally as of 2008, accounting for 30% to the vertebral column, and the front surface sits deep
to the sternum and costal cartilages.[7] Two of the great
1

1 STRUCTURE

Layers of the heart wall, including visceral and parietal pericardium.

epithelium, and covers heart chambers and valves. It is


continuous with the endothelium of the veins and arteries of the heart, and is joined to the myocardium with a
thin layer of connective tissue.[7] The endocardium, by
secreting endothelins, may also play a role in regulating
the contraction of the myocardium.[7]

The human heart is in the middle of the thorax, with its apex
pointing to the left.[12]

veins the venae cavae, and the great arteries, the aorta
and pulmonary artery, are attached to the upper part of
the heart, called the base, which is located at the level
of the third costal cartilage.[7] The lower tip of the heart,
the apex, lies just to the left of the sternum between the
junction of the fourth and fth ribs near their articulation
with the costal cartilages.[7] The right side of the heart is
deected forwards, and the left deected to the back.[7]
The heart is cone-shaped, with its base positioned upwards and tapering down to the apex.[7] A stethoscope can
be placed directly over the apex so that the heartbeats can
be counted. An adult heart has a mass of 250350 grams
(912 oz).[14] The heart is typically the size of a st: 12
cm (5 in) in length, 8 cm (3.5 in) wide, and 6 cm (2.5
in) in thickness.[7] Well-trained athletes can have much
larger hearts due to the eects of exercise on the heart
muscle, similar to the response of skeletal muscle.[7]

The swirling pattern of myocardium helps the heart pump eectively

The middle layer of the heart wall is the myocardium,


which is the cardiac muscle a layer of involuntary
striated muscle tissue surrounded by a framework of
collagen. The myocardium is also supplied with blood
1.1 Heart wall
vessels, and nerve bers by way of the epicardium that
help to regulate the heart rate.[7] Cardiac muscle tisMain article: Cardiac muscle
sue has autorhythmicity, the unique ability to initiate a
The heart wall is made up of three layers: the inner cardiac action potential at a xed rate spreading the imendocardium, middle myocardium and outer epicardium. pulse rapidly from cell to cell to trigger the contraction of
These are surrounded by a double-membraned sac called the entire heart. This autorhythmicity is still modulated
the pericardium.
by the endocrine and nervous systems.[7]
The innermost layer of the heart is called the endo- There are two types of cardiac muscle cell:
cardium. It is made up of a lining of simple squamous cardiomyocytes which have the ability to contract

1.2

Chambers

easily, and modied cardiomyocytes the pacemaker cells


of the conducting system. The cardiomyocytes make
up the bulk (99%) of cells in the atria and ventricles.
These contractile cells are connected by intercalated
discs which allow a rapid respond to impulses of action
potential from the pacemaker cells. The intercalated
discs allow the cells to act as a syncytium and enable the
contractions that pump blood through the heart and into
the major arteries.[7]

the atrioventricular valves and separated from the ventricles by the coronary sulcus. There is an ear-shaped structure in the upper right atrium called the right atrial appendage, or auricle, and another in the upper left atrium,
the left atrial appendage. The right atrium and the right
ventricle together are sometimes referred to as the right
heart and this sometimes includes the pulmonary artery.
Similarly, the left atrium and the left ventricle together
are sometimes referred to as the left heart. The ventricles
The pacemaker cells make up just (1% of cells) and form are separated by the anterior longitudinal sulcus and the
posterior interventricular sulcus.
the conduction system of the heart. They are generally much smaller than the contractile cells and have few The cardiac skeleton is made of dense connective tismyobrils which gives them limited contractibility. Their sue and this gives structure to the heart. It forms the
function is similar in many respects to neurons.[7]
atrioventricular septum which separates the atria from the
rings which serve as bases for
The cardiac muscle pattern is elegant and complex, as the ventricles, and the brous
[17]
the
four
heart
valves.
The
cardiac skeleton also promuscle cells swirl and spiral around the chambers of the
vides
an
important
boundary
in
the hearts electrical con[7]
heart. They form a gure 8 pattern around the atria and
duction
system
since
collagen
cannot
conduct electricity.
[7]
around the bases of the great vessels. Deeper ventricuThe
interatrial
septum
separates
the
atria
and the interlar muscles also form a gure 8 around the two ventricles
[7]
ventricular
septum
separates
the
ventricles.
The interand proceed toward the apex. More supercial layers of
ventricular
septum
is
much
thicker
than
the
interatrial
[7]
ventricular muscle wrap around both ventricles. This
septum,
since
the
ventricles
need
to
generate
greater
prescomplex swirling pattern allows the heart to pump blood
[7]
sure
when
they
contract.
[7]
more eectively than a simple linear pattern would.
As with skeletal muscles the heart can increase in size and
eciency with exercise.[7] Thus endurance athletes such 1.2.1 Valves
as marathon runners may have a heart that has increased
Main article: Heart valves
in size by up to 40%.[15]
The pericardium surrounds the heart. It consists of two
membranes: an inner serous membrane called the epicardium, and an outer brous membrane.[16] These enclose the pericardial cavity which contains the pericardial
uid that lubricates the surface of the heart.

1.2

Chambers

With the atria and major vessels removed, all four valves
are clearly visible.[7]
Superior
Vena Cava

Aorta

Pulmonary
Artery
Pulmonary
Vein

Right
Atrium

Left
Atrium
Mitral
Valve

Left
Ventricle

Pulmonary
Valve
Tricuspid
Valve

Heart being dissected showing right and left ventricles, from


above

Aortic
Valve

Right
Ventricle

Inferior Vena Cava

The heart, showing valves, arteries and veins. The white


arrows shows the normal direction of blood ow.
The heart has four chambers, two upper atria, the receivAll four heart valves lie along the same plane. The
ing chambers, and two lower ventricles, the discharging valves ensure unidirectional blood ow through the heart
chambers. The atria are connected to the ventricles by and prevent backow. Between the right atrium and the

1 STRUCTURE
1.2.2 Right heart

Frontal section showing papillary muscles attached to the tricuspid valve on the right and to the mitral valve on the left via chordae tendineae.[7]

right ventricle is the tricuspid valve. This consists of three


cusps (aps or leaets), made of endocardium reinforced
with additional connective tissue. Each of the three
valve-cusps is attached to several strands of connective
tissue, the chordae tendineae (tendinous cords), sometimes referred to as the heart strings. They are composed
of approximately 80 percent collagenous bers with the
remainder consisting of elastic bers and endothelium.
They connect each of the cusps to a papillary muscle that
extends from the lower ventricular surface. These muscles control the opening and closing of the valves. The
three papillary muscles in the right ventricle are called
the anterior, posterior, and septal muscles, which correspond to the three positions of the valve cusps.

The two major systemic veins, the superior and inferior


venae cavae, and the collection of veins that make up the
coronary sinus which drains the myocardium, empty into
the right atrium. The superior vena cava drains blood
from above the diaphragm and empties into the upper
back part of the right atrium. The inferior vena cava
drains the blood from below the diaphragm and empties
into the back part of the atrium below the opening for the
superior vena cava. Immediately above and to the middle
of the opening of the inferior vena cava is the opening of
the thin-walled coronary sinus.[7]
In the wall of the right atrium is an oval-shaped depression known as the fossa ovalis, which is a remnant of an
opening in the fetal heart known as the foramen ovale.
The foramen ovale allowed blood in the fetal heart to pass
directly from the right atrium to the left atrium, allowing some blood to bypass the pulmonary circuit. Within
seconds after birth, a ap of tissue known as the septum
primum that previously acted as a valve closes the foramen ovale and establishes the typical cardiac circulation
pattern.[7] Most of the internal surface of the right atrium
is smooth, the depression of the fossa ovalis is medial,
and the anterior surface has prominent ridges of pectinate
muscles, which are also present in the right atrial appendage.[7]

The atria receive venous blood on a nearly continuous basis, preventing venous ow from stopping while the ventricles are contracting. While most ventricular lling occurs while the atria are relaxed, they do demonstrate a
contractile phase when they actively pump blood into the
ventricles just prior to ventricular contraction. The right
Between the left atrium and left ventricle is the mitral atrium[7]is connected to the right ventricle by the tricuspid
valve, also known as the bicuspid valve due to its having valve.
two cusps, an anterior and a posterior medial cusp. These When the myocardium of the ventricle contracts, prescusps are also attached via chordae tendinae to two pap- sure within the ventricular chamber rises. Blood, like
illary muscles projecting from the ventricular wall.
any uid, ows from higher pressure to lower pressure
The tricuspid and the mitral valves are the atrioventric- areas, in this case, toward the pulmonary artery and the
atrium. To prevent any potential backow, the papillary
ular valves. During the relaxation phase of the cardiac
cycle, the papillary muscles are also relaxed and the ten- muscles also contract, generating tension on the chordae
tendineae. This prevents the aps of the valves from besion on the chordae tendineae is slight. However, as the
ventricle contracts, so do the papillary muscles. This cre- ing forced into the atria and regurgitation of the[7]blood
ates tension on the chordae tendineae, helping to hold the back into the atria during ventricular contraction.
cusps of the atrioventricular valves in place and prevent- The walls of the right ventricle are lined with trabeculae
carneae, ridges of cardiac muscle covered by endoing them from being blown back into the atria.[7]
The semilunar pulmonary valve is located at the base of cardium. In addition to these muscular ridges, a band of
the pulmonary artery. This has three cusps which are not cardiac muscle, also covered by endocardium, known as
attached to any papillary muscles. When the ventricle re- the moderator band reinforces the thin walls of the right
laxes blood ows back into the ventricle from the artery ventricle and plays a crucial role in cardiac conduction.
and this ow of blood lls the pocket-like valve, press- It arises from the lower part of the interventricular sepventricle
ing against the cusps which close to seal the valve. The tum and crosses the interior space of the right
[7]
to
connect
with
the
inferior
papillary
muscle.
semilunar aortic valve is at the base of the aorta and also
is not attached to papillary muscles. This too has three When the right ventricle contracts, it ejects blood into
cusps which close with the pressure of the blood owing the pulmonary artery, which branches into the left and
back from the aorta.[7]
right pulmonary arteries that carry it to each lung. The

5
upper surface of the right ventricle begins to taper as it
approaches the pulmonary artery. At the base of the pulmonary artery is the pulmonary semilunar valve that prevents backow from the pulmonary artery.[7]
1.2.3

Left heart

branches of these arteries anastomose, which in other


parts of the body serve to divert blood due to a blockage.
In the heart these are very small and cannot form other
interconnections with the result that a coronary artery
blockage can cause a myocardial infarction and with it,
tissue damage.[7]
The great cardiac vein receives the major branches of the
posterior, middle, and small cardiac veins and drains into
the coronary sinus, a large vein that empties into the right
atrium. The anterior cardiac veins drain the front of the
right ventricle and drain directly into the right atrium.[7]

After gas exchange in the pulmonary capillaries, blood


high in oxygen returns to the left atrium via one of the four
pulmonary veins. Only the left atrial appendage contains
pectinate muscles. Blood ows nearly continuously from
the pulmonary veins back into the atrium, which acts as
the receiving chamber, and from here through an opening
into the left ventricle. Most blood ows passively into the 2 Development
heart while both the atria and ventricles are relaxed, but
toward the end of the ventricular relaxation period, the Main articles: Heart development and Human embryoleft atrium will contract, pumping blood into the ventri- genesis
cle. This atrial contraction accounts for approximately 20 The heart is the rst functional organ to develop and
percent of ventricular lling. The left atrium is connected
to the left ventricle by the mitral valve.[7]
Although both sides of the heart will pump the same
amount of blood, the muscular layer is much thicker in
the left ventricle compared to the right, due to the greater
force needed here. Like the right ventricle, the left also
has trabeculae carneae, but there is no moderator band.
The left ventricle is the major pumping chamber for the
systemic circuit; it ejects blood into the aorta through the
aortic semilunar valve.[7]

1.3

Coronary circulation

Superior vena cava


Aorta

Left pulmonary artery


Left pulmonary veins
Left coronary artery

Right atrium
Right coronary
artery
Posterior
descending artery
Right
marginal artery

Left circumflex artery


Left marginal artery
Left anterior descending
(or interventricular) artery
diagonal branch

Development of the human heart during the rst eight weeks


(top), and the formation of the heart chambers (bottom).[7]

starts to beat and pump blood at about three weeks into


embryogenesis. This early start is crucial for subsequent
embryonic and prenatal development.

Cardiomyocytes, like all other cells, need to be supplied


with oxygen, nutrients and a way of removing metabolic
wastes. This is achieved by the coronary circulation. The
coronary circulation cycles in peaks and troughs relating
to the heart muscle relaxing or contracting.[7]

The heart derives from splanchnopleuric mesenchyme in


the neural plate which forms the cardiogenic region. Two
endocardial tubes form here that fuse to form a primitive
heart tube known as the tubular heart.[18] Between the
third and fourth week, the heart tube lengthens, and begins to fold to form an S-shape within the pericardium.
This places the chambers and major vessels into the correct alignment for the developed heart. Further development will include the septa and valves formation and
remodelling of the heart chambers. By the end of the
fth week the septa are complete and the heart valves are
completed by the ninth week.[7]

Coronary arteries supply oxygen-rich blood to the heart


and the coronary veins remove the deoxygenated blood.
There is a left and a right coronary artery supplying the
left and right hearts respectively, and the septa. Smaller

The embryonic heart begins beating at around 22 days


after conception (5 weeks after the last normal menstrual period, LMP). It starts to beat at a rate near to the
mothers which is about 7580 beats per minute (bpm).

Right ventricle

Left ventricle

Arterial supply to the heart (red), with other areas labelled (blue).

Main article: Coronary circulation

3 PHYSIOLOGY

The embryonic heart rate then accelerates and reaches a


peak rate of 165185 bpm early in the early 7th week
(early 9th week after the LMP).[19][20][21] After 9 weeks
(start of the fetal stage) it starts to decelerate, slowing to
around 145 (25) bpm at birth. There is no dierence in
female and male heart rates before birth.[22]

Physiology

Main article: Cardiac physiology

transports oxygen to the body and returns carbon dioxide


and relatively deoxygenated blood to the heart for transfer
to the lungs.[7]
The right heart collects deoxygenated blood from two
large veins, the superior and inferior venae cavae. The
blood collects in the right atrium and is pumped through
the tricuspid valve into the right ventricle, where it is
pumped into the pulmonary artery through the pulmonary
valve. Here the blood enters the pulmonary circulation
where carbon dioxide can be exchanged for oxygen in
the lungs. This happens through the passive process of
diusion.
In the left heart, oxygenated blood is returned to the
left atrium via the pulmonary vein. It is then pumped
into the left ventricle through the bicuspid valve and into
the aorta through the mitral valve for systemic circulation. The aorta is a large artery that branches into many
smaller arteries, arterioles, and ultimately capillaries. In
the capillaries, oxygen and nutrients from blood are supplied to body cells for metabolism, and exchanged for carbon dioxide and waste products[7]
3.1.1 Cardiac cycle

Blood ow through the heart from the Khan academy

Main articles: Cardiac cycle, systole and diastole


The cardiac cycle refers to a complete heartbeat which in-

3.1

Blood ow

The cardiac cycle as correlated to the ECG


Blood ow through the valves

The heart functions as a pump in the circulatory system to


provide a continuous circulation of blood throughout the
body. This circulation consists of the systemic circulation
to and from the body and the pulmonary circulation to
and from the lungs. Blood in the pulmonary circulation
exchanges carbon dioxide for oxygen in the lungs through
the process of respiration. The systemic circulation then

cludes systole and diastole and the intervening pause. The


cycle begins with contraction of the atria and ends with
relaxation of the ventricles. Systole is when the ventricles
of the heart contract to pump blood to the body. Diastole is when the ventricles relax and ll with blood. The
atria and ventricles work in concert, so in systole when
the ventricles are contracting, the atria are relaxed and
collecting blood. When the ventricles are relaxed in di-

3.2

Electrical conduction

astole, the atria contract to pump blood to the ventricles. Preload refers to how much blood is in the ventricles at the
This coordination ensures blood is pumped eciently to end of diastole, at their most full. A main factor is how
the body.[7]
long it takes the ventricles to llif the ventricles conthen there is less time to ll and the preload
At the beginning of the cardiac cycle, in early diastole, tract faster, [7]
will
be
less.
Preload can also be aected by a persons
both the atria and ventricles are relaxed. Since blood
hydration
status.
. It is important because of the Frankmoves from areas of high pressure to areas of low presStarling
mechanism.
This states that the force of contracsure, when the chambers are relaxed, blood will ow into
tion
is
directly
proportional
to the initial length of musthe atria (through the coronary sinus and the pulmonary
cle ber. This means that a ventricle will contract more
veins). As the atria begin to ll, the pressure will rise
[7]
so that the blood will move from the atria into the ven- forcefully, the more it is stretched.
tricles. In late diastole the atria contract pumping more
blood into the ventricles. This causes a rise in pressure
in the ventricles, and in ventricular systole blood will be
pumped into the pulmonary artery.

Afterload, or how much blood is left in the ventricles after


systole, is inuenced by the resistance of the vascular system. This tension is called afterload. It can be inuenced
by narrowing of the heart valves (stenosis) or contraction
[7]
When the atrioventricular valves (tricuspid and mitral) are or relaxation of the peripheral blood vessels.
open, during blood ow to the ventricles, the semilunar The strength of heart muscle contractions controls the
valves are closed to prevent backow into the ventricles. stroke volume. This can be inuenced positively or negWhen the ventricular pressure is greater than the atrial atively by agents termed inotropes. These can be either
pressure the tricuspid and mitral valves will shut. When conditions or drugs. Positive inotropes that cause stronger
the ventricles contract the pressure forces the semilunar contractions include high blood calcium and drugs such
aortic and pulmonary valves open. As the ventricles relax as Digoxin, which will act to stimulate the sympathetic
the semilunar valves will close in response to decreased nerves in the ght-or-ight response. Negative inotropes
pressure.
causing weaker contractions include high blood potassium, hypoxia, acidosis, and drugs such as beta blockers and calcium channel blockers. These act on the
3.1.2 Cardiac output
parasympathetic nervous system via the vagus nerve.
Main article: Cardiac output
Cardiac output (CO) is a measurement of the amount of 3.2

Electrical conduction

Main article: Electrical conduction system of the heart


The normal rhythmical heart beat, called sinus rhythm, is

The x-axis reects time with a recording of the heart sounds. The
y-axis represents pressure.[7]

blood pumped by each ventricle in one minute. To calculate this, multiply the amount pumped out by each ventricle, the stroke volume (SV), by the heart rate (HR), in
beats per minute.[7] Cardiac output can be represented by
the equation: CO = HR x SV[7] The average cardiac output, using an average SV of about 70mL, is 5.25 L/min,
with a range of 4.08.0 L/min.[7] The stroke volume is
normally measured using an echocardiogram and can be Transmission of a cardiac action potential through the hearts
inuenced by the size of the heart, physical and mental conduction system
condition of the individual, sex, contractility, duration of
contraction, preload and afterload.[7]
established by the sinoatrial node, the hearts pacemaker.

3 PHYSIOLOGY

Here an electrical signal is created that travels through the


heart, causing the heart muscle to contract.
The sinoatrial node is found in the coronary sinus of
the right atrium.[7] The electrical signal generated by the
sinoatrial node travels through the right atrium in a radial way that is not completely understood. It travels to
the left atrium via Bachmanns bundle, such that both left
and right atrium contract together.[23][24][25] The signal
then travels to the atrioventricular node. This is found at
the bottom of the right atrium in the atrioventricular septumthe boundary between the right atrium and the left
ventricle. The septum is part of the cardiac skeleton, tissue within the heart that the electrical signal cannot pass
through, which forces the signal to pass through the atrioventricular node only.[7] The signal then travels along the
Bundle of His to left and right bundle branches through
to the ventricles of the heart. In the ventricles the signal
is carried by specialized tissue called the Purkinje bers
which then transmit the electric charge to the cardiac
muscle.[26]
Sinoatrial node

Atrioventricular
node

Bachmann's
bundle

His bundle

Left posterior
bundle
Right bundle

Conduction system of the heart

3.3

Heart rate

Main article: Heart rate

Purkinje
fibers

The prepotential is due to a slow inux of sodium ions until the


threshold is reached followed by a rapid depolarization and repolarization. The prepotential accounts for the membrane reaching
threshold and initiates the spontaneous depolarization and contraction of the cell; there is no resting potential.[7]

When the sinoatrial cells are resting, they have a negative charge on their membranes. However a rapid inux of sodium ions causes the membranes charge to
become positive. This is called depolarisation and occurs spontaneously.[7] Once the cell has a suciently high
charge, the sodium channels close and calcium ions then
begin to enter the cell, shortly after which potassium begins to leave it. All the ions travel through ion channels in
the membrane of the sinoatrial cells. The potassium and
calcium only start to move out of and into the cell once it
has a suciently high charge, and so are called voltagegated. Shortly after this, the calcium channels close and
potassium channels open, allowing potassium to leave the
cell. This causes the cell to have a negative resting charge
and is called repolarization. When the membrane potential reaches approximately 60 mV, the potassium channels close and the process may begin again.[7]
The ions move from areas where they are concentrated to
where they are not. For this reason sodium moves into
the cell from outside, and potassium moves from within
the cell to outside the cell. Calcium also plays a critical
role. Their inux through slow channels means that the
sinoatrial cells have a prolonged pleateau phase when
they have a positive charge. A part of this is called the
absolute refractory period. Calcium ions also combine
with the regulatory protein troponin C in the troponin
complex to enable contraction of the cardiac muscle, and
separate from the protein to allow relaxation.[27]

The resting heart rate of a newborn can be 120 beats per


minute (bpm) and this gradually decreases until maturity
and then gradually increases again with age. The adult
resting heart rate ranges from 60 to 100 bpm. Exercise
and tness levels, age and basal metabolic rate can all affect the heart rate. An athletes heart rate can be lower
than 60bpm. During exercise the rate can be 150bpm 3.3.2 Inuences
with maximum rates reaching from 200 and 220 bpm.[7]
The normal sinus rhythm of the heart, giving the resting
heart rate, is inuenced by the autonomic nervous sys3.3.1 Creation
tem through sympathetic and parasympathetic nerves.[28]
These arise from two paired cardiovascular centres in
The sinoatrial node create and sustains its own rhythm. the medulla oblongata.The vagus nerve of the parasymCells in the sinoatrial node do this by creating an action pathetic nervous system acts to decrease the heart rate,
potential. The cardiac action potential is created by the and nerves from the sympathetic trunk act to increase
movement of specic electrolytes into and out of the the heart rate. These come together in the cardiac plexus
pacemaker cells. The action potential then spreads to near the base of the heart. Without parasympathetic input
which normally predominates, the sinoatrial node would
nearby cells.

3.4

Heart sounds

9
of receptors including proprioreceptors, baroreceptors,
and chemoreceptors, plus stimuli from the limbic system.
Through a series of reexes these help regulate and sustain blood ow. For example, increased physical activity
results in increased rates of ring by various proprioreceptors located in muscles, joint capsules, and tendons.
With increased rates of ring, the parasympathetic stimulation may decrease or sympathetic stimulation may increase as needed in order to increase blood ow.[7]
Similarly, baroreceptors are stretch receptors located in
the aortic sinus, carotid bodies, the venae cavae, and other
locations, including pulmonary vessels and the right side
of the heart itself. Rates of ring from the baroreceptors
represent blood pressure, level of physical activity, and
the relative distribution of blood. The cardiac centers
monitor baroreceptor ring to maintain cardiac homeostasis, a mechanism called the baroreceptor reex. With
increased pressure and stretch, the rate of baroreceptor
ring increases, and the cardiac centers decrease sympathetic stimulation and increase parasympathetic stimulation. As pressure and stretch decrease, the rate of
baroreceptor ring decreases, and the cardiac centers increase sympathetic stimulation and decrease parasympathetic stimulation.[7]
There is a similar reex, called the atrial reex or
Bainbridge reex, associated with varying rates of blood
ow to the atria. Increased venous return stretches the
walls of the atria where specialized baroreceptors are located. However, as the atrial baroreceptors increase their
rate of ring and as they stretch due to the increased blood
pressure, the cardiac center responds by increasing sympathetic stimulation and inhibiting parasympathetic stimulation to increase HR. The opposite is also true.[7]

Autonomic innervation of the heart

generate a heart rate of about 100 bpm.[7]


The nerves from the sympathetic trunk emerge through
the T1-T4 thoracic ganglia and travel to both the sinoatrial and atrioventricular nodes, as well as to the atria and
ventricles. The ventricles are more richly innervated by
sympathetic bers than parasympathetic bers. Sympathetic stimulation causes the release of the neurotransmitter norepinephrine (also known as noradrenaline) at
the neuromuscular junction of the cardiac nerves. This
shortens the repolarization period, thus speeding the rate
of depolarization and contraction, which results in an increased heart rate. It opens chemical or ligand-gated
sodium and calcium ion channels, allowing an inux of
positively charged ions.[7] Norepinephrine binds to the
beta1 receptor. High blood pressure medications are
used to block these receptors and so reduce the heart
rate.[7]

In addition to the autonomic nervous system, other factors can impact on this. These include epinephrine, norepinephrine, and thyroid hormones; levels of various ions
including calcium, potassium, and sodium; body temperature; hypoxia; and pH balance. Factors that increase
the heart rate can include release of norepinephrine,
hypoxemia, low blood pressure and dehydration, a strong
emotional response, a higher body temperature, and
metabolic and hormonal factors such as a low potassium
or sodium level or stimulus from thyroid hormones.[7] Decreased body temperature, relaxation, and metabolic factors can also contribute to a decrease in heart rate.[7]

3.4 Heart sounds


Main article: Heart sounds

One of the simplest methods of assessing the hearts condition is to listen to it using a stethoscope.[7] In a healthy
heart, there are only two audible heart sounds, called S1
and S2. The rst heart sound S1, is the sound created
by the closing of the atrioventricular valves during venThe cardiovascular centres receive input from a series tricular contraction and is normally described as lub.

10

4 CLINICAL SIGNIFICANCE
listening for heart murmurs.

Atherosclerosis is a condition aecting the circulatory


system. If the coronary arteries are aected angina
pectoris may result or at worse a heart attack.
3D echocardiogram showing the mitral valve (right), tricuspid
and mitral valves (top left) and aortic valve (top right).
The closure of the heart valves causes the heart sounds.

The second heart sound, S2, is the sound of the semilunar valves closing during ventricular diastole and is described as dub.[7] Each sound consists of two components, reecting the slight dierence in time as the two
valves close.[29] S2 may split into two distinct sounds, either as a result of inspiration or dierent valvular or cardiac problems.[29] Additional heart sounds may also be
present and these give rise to gallop rhythms. A third
heart sound, S3 usually indicates an increase in ventricular blood volume. A fourth heart sound S4 is referred to
as an atrial gallop and is produced by the sound of blood
being forced into a sti ventricle. The combined presence of S3 and S4 give a quadruple gallop.[7]
Heart murmurs are abnormal heart sounds which can
be either pathological or benign.[30] One example of
a murmur is Stills murmur, which presents a musical
sound in children, has no symptoms and disappears in
adolescence.[31]
A dierent type of sound, a pericardial friction rub can
be heard in cases of pericarditis where the inamed membranes can rub together.[32]

Clinical signicance

Being such a complex organ the heart is prone to several cardiovascular diseases some becoming more prevalent with ageing.[33] Heart disease is a major cause of
death, accounting for an average of 30% of all deaths in
2008, globally.[9] This rate varies from a lower 28% to a
high 40% in high-income countries.[10] Doctors that specialise in the heart are called cardiologists. Many other
medical professionals are involved in treating diseases of
the heart, including doctors such as general practitioners,
cardiothoracic surgeons and intensivists, and allied health
practitioners including physiotherapists and dieticians.
Obesity, high blood pressure, and high cholesterol can
all increase the risk of developing heart disease. However, half the number of heart attacks occur in people
with normal cholesterol levels. It is generally accepted
that factors such as exercise or the lack of it, good or
poor diet, and overall well-being (including emotional),
aect heart health.[34][35][36][37] Exercise results in the
addition of protein myolaments and this can result in
hypertrophy where the size of individual cells are increased but not their number.[7] This is a condition known
as athletic heart syndrome. The hearts of athletes can
pump more eciently at lower heart rates. However,
enlarged hearts can have a pathological cause such as
hypertrophic cardiomyopathy, which can result in a heart
of 1000 g (2 lb) in mass.[7] The cause of an abnormally
enlarged heart muscle is unknown, but the condition is
often undiagnosed and can cause sudden death in young
athletes.[7]
Coronary artery disease is also known as ischemic heart
disease, and atherosclerotic disease and is the most common form of heart disease. The underlying mechanism of
this disease is atherosclerosisa build-up of plaque along
the inner walls of the arteries which narrows them, reducing the blood ow to the heart. It is the leading cause
of heart attacks and the most common cause of death,
globally.[38] It is also the main cause of angina.

Cardiomyopathy and most commonly dilated cardiomyopathy, is a noticeable deterioration of the heart muscles
ability to contract, which can lead to heart failure.[39][40]
Other common causes of heart failure (which can also be
The stethoscope is used for auscultation of the heart, congestive), are heart attacks, valve disorders and high
and is one of the most iconic symbols for medicine. blood pressure. This happens when the heart is pumpA number of diseases can be detected primarily by

4.2

Assessment

11

ing insuciently and cannot meet the bodys blood ow


demands.[41] Because the heart is a double pump, each
side can fail independently of the other, resulting in heart
failure of the right heart or the left heart, either of which
through causing strain in the other side can result in the
failure of the whole heart. Congestive heart failure results
in blood backing up in the systemic circulation. Edema
(swelling) of the feet, ankles and ngers is the most noticeable symptom. Pulmonary congestion results from
left heart failure. The right side of the heart continues
to propel blood to the lungs, but the left side is unable to
pump the returning blood into the systemic circulation.
As blood vessels within the lungs become swollen with
blood, the pressure within them increases, and uid leaks
from the circulation into the lung tissue. This pleural effusion causes pulmonary edema. If untreated, the person will suocate because they are drowning in their own
blood.[42]

The cardiac examination includes inspection, palpation


and auscultation.

Heart murmurs are abnormal heart sounds which can


be either pathological or benign and there are several
kinds.[30] Murmurs are graded by volume, from 1) the
quietest, to 6) the loudest, and evaluated by their relationship to the heart sounds and position in the cardiac
cycle.[29] Phonocardiograms can record these sounds.[7]
Murmurs can result from valvular heart diseases due
to narrowing (stenosis), regurgitation or insuciency of
any of the main heart valves but they can also result
from a number of other disorders, including atrial and
ventricular septal defects.[29]

Cardiac cycle shown against ECG

Abnormalities in the sinus rhythm can prevent the heart


from eectively pumping blood and cause both atrial and
ventricular brillation.[42] Examples of cardiac dysrhythmias are a very rapid heart rate (tachycardia) and a very
slow heart rate (brachycardia). Tachycardia is generally
dened as a heart rate faster than 100 beats per minute,
and bradycardia as a heart rate slower than 60. Asystole
is the cessation of heart rhythm which results in cardiac
arrest.
Cardiac tamponade, also known as pericardial tamponade, is the condition of an abnormal build-up of uid
in the pericardium which can adversely aect the function of the heart. The uid can be removed from the
pericardial sac using a syringe in a procedure called
pericardiocentesis.
Carditis is inammation of the heart; this can be specic
to regions as in pericarditis, myocarditis, and endocarditis
or it can be of the whole heart known as pancarditis.

4.1

Diagnosis

4.2

Assessment

4.2.1

Examination

Main article: Cardiac examination

4.2.2 Electrocardiogram
Main article: Electrocardiography
Using surface electrodes on the body, it is possible to

record the complex electrical activity of the heart. This


tracing of the electrical signal is the electrocardiogram
(ECG) or (EKG). An ECG clearly shows normal and abnormal heart function and is an indispensable diagnostic
tool.
There are ve prominent points on the ECG: the P wave
(atrial depolarisation), the QRS complex (atrial repolarisation and ventricular depolarisation) and the T wave
(ventricular repolarisation).[7]
4.2.3 Imaging
Main article: Cardiac imaging
Several imaging methods can be used to assess the
anatomy and function of the heart, including ultrasound,
angiography, PET, CT and MRI. Ultrasound of the
heart is called echocardiography. It is used to measure
the hearts function, assess for disease of the valves of
the heart, and look for any anatomical abnormalities.
Echocardiography can be conducted by a probe on the
chest (transthoracic) or by a probe in the esophagus
(transoesophageal). A typical echocardiography report
will include information about the volumes at the end
of systole and diastole, how wide the valves are (checking for stenosis), whether there is any backow of blood
through the valves (regurgitation), and an ejection fraction, which describes how much blood is ejected from
the left and right ventricles after systole. Ejection fractions range from approximately 55 to 70 percent, with a
mean of 58 percent.[7]
4.2.4 Stress tests
Main article: Cardiac stress test

12

5 HISTORY

A cardiac stress test uses exercise or drugs to stimulate of the Hippocratean school around the 4th century BC,
the heart and provoke a measurable response to the stress although their function was not fully understood. On
in order to gauge the hearts eectiveness.
dissection, arteries are typically empty of blood because
blood pools in the veins after death. It was subsequently
assumed they were lled with air and served to transport
4.3 Treatment
air around the body.
Philosophers distinguished veins from arteries, but
Angiogenesis represents a therapeutic target for cardiothought the pulse was a property of arteries. Erasistratos
vascular disease.[43]
observed that arteries cut during life bleed. He ascribed
Debrillation is used to treat serious arrhythmias. the fact to the phenomenon that air escaping from an
Articial pacemakers used to regulate the heartbeat can artery is replaced with blood which entered by very small
also incorporate a debrillator.
vessels between veins and arteries. Thus he apparently
postulated capillaries, but with reversed ow of blood.

4.4

Surgery

Main articles: Coronary artery bypass surgery and


Coronary stent
Coronary artery bypass surgery to improve the blood supply to the heart is often the only treatment option for coronary heart disease.
Heart valve repair or valve replacement are options for
valvular heart disease.

5
5.1

History
Ancient

The Greek physician Galen (2nd century AD) knew blood


vessels carried blood and identied venous (dark red) and
arterial (brighter and thinner) blood, each with distinct
and separate functions. Growth and energy were derived
from venous blood created in the liver from chyle, while
arterial blood gave vitality by containing pneuma (air) and
originated in the heart. Blood owed from both creating
organs to all parts of the body, where it was consumed
and there was no return of blood to the heart or liver.
The heart did not pump blood around, the hearts motion
sucked blood in during diastole and the blood moved by
the pulsation of the arteries themselves.
Galen believed the arterial blood was created by venous
blood passing from the left ventricle to the right through
'pores in the interventricular septum, while air passed
from the lungs via the pulmonary artery to the left side
of the heart. As the arterial blood was created, sooty
vapors were created and passed to the lungs, also via the
pulmonary artery, to be exhaled.

5.2 Pre-modern
The earliest descriptions of the coronary and pulmonary
circulation systems can be found in the Commentary on
Anatomy in Avicennas Canon, published in 1242 by Ibn
al-Nas.[44] In his manuscript, al-Nas wrote that blood
passes through the pulmonary circulation instead of moving from the right to the left ventricle as previously believed by Galen.[45] His work was later translated into
Latin by Andrea Alpago.[46]
In Europe, the teachings of Galen continued to dominate
the academic community and his doctrines were adopted
as the ocial canon of the Church. Andreas Vesalius
questioned some of Galens beliefs of the heart in De humani corporis fabrica (1543), but his magnum opus was
interpreted as a challenge to the authorities and he was
Heart and its blood vessels, by Leonardo da Vinci, 15th century subjected to a number of attacks.[47] Michael Servetus
wrote in Christianismi Restitutio (1553) that blood ows
The valves of the heart were discovered by a physician from one side of the heart to the other via the lungs.[47]

13

5.3

Modern

6 Society and culture


Further information: Sacred Heart, Heart symbol and
Blood Cultural and religious beliefs

The breakthrough came with the publication of De Motu


Cordis (1628) by the English physician William Harvey.
6.1 Symbolism
Harveys book completely describes the systemic circulation and the mechanical force of the heart, leading to an
overhaul of the Galenic doctrines.[48] Otto Frank (1865
1944) was a German physiologist; among his many published works are detailed studies of this important heart
relationship. Ernest Starling (18661927) was an important English physiologist who also studied the heart.
Although they worked largely independently, their combined eorts and similar conclusions have been recogLetter of the Georgian script is often used as a heart
nized in the name "FrankStarling mechanism.[7]
symbol.
Although Purkinje bers and the bundle of His were discovered as early as the 19th century, their specic role
in the electrical conduction system of the heart remained
unknown until Sunao Tawara published his monograph,
titled Das Reizleitungssystem des Sugetierherzens, in
1906. Tawaras discovery of the atrioventricular node
prompted Arthur Keith and Martin Flack to look for similar structures in the heart, leading to their discovery of The seal script glyph for heart (Middle Chinese sim)
the sinoatrial node several months later. These structures form the anatomical basis of the electrocardiogram, As one of the vital organs, the heart was long identiwhose inventor, Willem Einthoven, was awarded the No- ed as the center of the entire body, the seat of life, or
bel Prize in Medicine or Physiology in 1924.[49]
emotion, or reason, will, intellect, purpose or the mind.
The rst successful heart transplantation was performed Thus, in the Hebrew Bible, the word for heart lebab
in 1967 by the South African surgeon Christiaan Barnard is used in these meanings (paralleling the use of
at Groote Schuur Hospital in Cape Town. This marked "diaphragm" in Homeric Greek).
an important milestone in cardiac surgery, capturing the
attention of both the medical profession and the world
at large. However, long-term survival rates of patients
were initially very low. Louis Washkansky, the rst recipient of a donated heart, died 18 days after the operation
while other patients did not survive for more than a few
weeks.[50] The American surgeon Norman Shumway has
been credited for his eorts to improve transplantation
techniques, along with pioneers Richard Lower, Vladimir
Demikhov and Adrian Kantrowitz. As of March 2000,
more than 55,000 heart transplantations have been performed worldwide.[51]
By the middle of the 20th century, heart disease had surpassed infectious disease as the leading cause of death in
the United States, and it is currently the leading cause of
deaths worldwide. Since 1948, the ongoing Framingham
Heart Study has shed light on the eects of various inuences on the heart, including diet, exercise, and common
medications such as aspirin. Although the introduction of
ACE inhibitors and beta blockers has improved the management of chronic heart failure, the disease continues
to be an enormous medical and societal burden, with 30
to 40% of patients dying within a year of receiving the
diagnosis.[52]

An important part of the concept of the soul in Ancient


Egyptian religion was thought to be the heart, or ib. The
ib or metaphysical heart was believed to be formed from
one drop of blood from the childs mothers heart, taken
at conception.[53] To ancient Egyptians, the heart was the
seat of emotion, thought, will, and intention. This is evidenced by Egyptian expressions which incorporate the
word ib, such as Awt-ib for happiness (literally, wideness of heart), Xak-ib for estranged (literally, truncated of heart). In Egyptian religion, the heart was the
key to the afterlife. It was conceived as surviving death in
the nether world, where it gave evidence for, or against, its
possessor. It was thought that the heart was examined by
Anubis and the deities during the Weighing of the Heart
ceremony. If the heart weighed more than the feather
of Maat, it was immediately consumed by the monster
Ammit.
The Chinese character for heart, , derives from a comparatively realistic depiction of a heart (indicating the
heart chambers) in seal script. The Chinese word xn also
takes the metaphorical meanings of mind, intelligence,
soul, or center, core. In Chinese medicine, the heart
is seen as the center of shn spirit, soul, consciousness.

14

The Sanskrit word for heart, hRd (), dates at least as


far back as the Rigveda and is a cognate of the word for
heart in Greek, Latin, and English. The same word is
used to mean mind or soul depending on the context.
Many classical philosophers and scientists, including
Aristotle, considered the heart the seat of thought, reason,
or emotion, often disregarding the brain as contributing
to those functions.[54] The identication of the heart as
the seat of emotions in particular is due to the Roman
physician Galen, who also located the seat of the passions
in the liver, and the seat of reason in the brain.[55] However these emotional properties of the heart were later
discovered to be solely centered in the brain. This tradition inuenced the development of the medieval Christian
devotion to the Sacred Heart of Jesus and the Immaculate
Heart of Mary.

OTHER ANIMALS

sausage. The Balinese oret is a sort of blood sausage made


with pig heart and blood. A French recipe for cur de
porc l'orange is made of braised heart with an orange
sauce.

7 Other animals
Further information: Circulatory system

The structure of the heart can vary among the dierent


animal species. Cephalopods have two gill hearts also
known as branchial hearts and one systemic heart. The
vertebrate heart lies in the front (ventral) part of the body
cavity, dorsal to the gut. It is always surrounded by a
pericardium, which is usually a distinct structure, but may
The idiomatic expression of pierced or broken hearts be continuous with the peritoneum in jawless and cartiultimately derive from devotional Christianity, where the laginous sh.
hearts of Mary or Jesus are depicted as suering various
The SA node is found in all amniotes but not in more
tortures (symbolizing the pain suered by Christ for the
primitive vertebrates. In these animals, the muscles of
sins of the world, and the pain of Mary at the crucixthe heart are relatively continuous and the sinus venoion of her son, respectively), but from an early time the
sus coordinates the beat which passes in a wave through
metaphor was transferred to unfullled romantic love, in
the remaining chambers. Indeed, since the sinus venolate medieval literature dealing with the ideals of courtly
sus is incorporated into the right atrium in amniotes, it is
love. The notion of "Cupid's arrows is ancient, due to
likely homologous with the SA node. In teleosts, with
Ovid, but while Ovid describes Cupid as wounding his
their vestigial sinus venosus, the main centre of coorvictims with his arrows, it is not made explicit that it is
dination is, instead, in the atrium. The rate of heartthe heart that is wounded. The familiar iconography of
beat varies enormously between dierent species, rangCupid shooting little heart symbols is Baroque.
ing from around 20 beats per minute in codsh to around
600 in hummingbirds[64] and up to 1200 bpm in the rubythroated hummingbird.[65]

6.2

Food

Animal hearts are widely consumed as food. As they are 7.1 Double circulatory systems
almost entirely muscle, they are high in protein. They are
often included in dishes with other oal, for example in In the heart of lungsh, the septum extends part-way into
the pan-Ottoman kokoretsi.
the ventricle. This allows for some degree of separation
Chicken hearts are considered to be giblets, and are of- between the de-oxygenated bloodstream destined for the
ten grilled on skewers: Japanese hto yakitori, Brazilian lungs and the oxygenated stream that is delivered to the
churrasco de corao, Indonesian chicken heart satay.[56] rest of the body. The absence of such a division in living
They can also be pan-fried, as in Jerusalem mixed grill. In amphibian species may be partly due to the amount of
Egyptian cuisine, they can be used, nely chopped, as part respiration that occurs through the skin; thus, the blood
of stung for chicken.[57] Many recipes combined them returned to the heart through the vena cavae is already
with other giblets, such as the Mexican pollo en menuden- partially oxygenated. As a result, there may be less need
for a ner division between the two bloodstreams than in
cias[58] and the Russian ragu iz kurinyikh potrokhov.[59]
lungsh or other tetrapods. Nonetheless, in at least some
The hearts of beef, pork, and mutton can generally be in- species of amphibian, the spongy nature of the ventricle
terchanged in recipes. As heart is a hard-working muscle, does seem to maintain more of a separation between the
it makes for rm and rather dry meat,[60] so is generally bloodstreams. Also, the original valves of the conus arslow-cooked. Another way of dealing with toughness is teriosus have been replaced by a spiral valve that divides
to julienne the meat, as in Chinese stir-fried heart.[61]
it into two parallel parts, thereby helping to keep the two
Beef heart may be grilled or braised.[60] In the Peruvian bloodstreams separate.[64]
anticuchos de corazn, barbecued beef hearts are grilled Adult amphibians and most reptiles have a double circulaafter being tenderized through long marination in a spice tory system but the heart is not separated into two pumps.
and vinegar mixture. An Australian recipe for mock The development of the double system is necessitated by
goose is actually braised stued beef heart.[62]
the presence of lungs which deliver oxygenated blood diPig heart is stewed, poached, braised,[63] or made into rectly to the heart.

7.3

Fish

15

7.3 Fish
Main article: Fish anatomy Heart
Primitive sh have a four-chambered heart, but the

A cross section of a three chambered adult amphibian heart, note


the single ventricle. The purple regions represent areas where
mixing of oxygenated and de-oxygenated blood occurs.
1. Pulmonary Vein
2. Left Atrium
3. Right Atrium
4. Ventricle
5. Conus arteriosus
6. Sinus venosus

In amphibians, the atrium is divided into two chambers


by a muscular septum but there is only one ventricle. The
sinus venosus, which remains large, connects only to the
right atrium and receives blood from the venae cavae,
with the pulmonary vein by-passing it to enter the left
atrium.
The heart of most reptiles is similar in structure to that of
lungsh but the septum is generally much larger. This divides the ventricle into two halves but the septum does not
reach the whole length of the heart and there is a considerable gap near the pulmonary artery and aorta openings.
In most reptilian species, there appears to be little, if any,
mixing between the bloodstreams, so the aorta receives,
essentially, only oxygenated blood.[64]

7.2

The fully divided heart

Archosaurs (crocodilians and birds) and mammals show


complete separation of the heart into two pumps for a
total of four heart chambers; it is thought that the fourchambered heart of archosaurs evolved independently
from that of mammals. In crocodilians, there is a small
opening, the foramen of Panizza, at the base of the arterial trunks and there is some degree of mixing between the blood in each side of the heart, during a dive
underwater;[66][67] thus, only in birds and mammals are
the two streams of blood those to the pulmonary and
systemic circulations permanently kept entirely separate by a physical barrier.[64]

Blood ow through the sh heart: sinus venosus, atrium,


ventricle, and outow tract

chambers are arranged sequentially so that this primitive heart is quite unlike the four-chambered hearts of
mammals and birds. The rst chamber is the sinus venosus, which collects deoxygenated blood, from the body,
through the hepatic and cardinal veins. From here, blood
ows into the atrium and then to the powerful muscular ventricle where the main pumping action will take
place. The fourth and nal chamber is the conus arteriosus which contains several valves and sends blood to
the ventral aorta. The ventral aorta delivers blood to the
gills where it is oxygenated and ows, through the dorsal
aorta, into the rest of the body. (In tetrapods, the ventral
aorta has divided in two; one half forms the ascending
aorta, while the other forms the pulmonary artery).[64]
In the adult sh, the four chambers are not arranged in a
straight row but, instead form an S-shape with the latter
two chambers lying above the former two. This relatively
simpler pattern is found in cartilaginous sh and in the
ray-nned sh. In teleosts, the conus arteriosus is very
small and can more accurately be described as part of
the aorta rather than of the heart proper. The conus arteriosus is not present in any amniotes, presumably having been absorbed into the ventricles over the course of
evolution. Similarly, while the sinus venosus is present
as a vestigial structure in some reptiles and birds, it is
otherwise absorbed into the right atrium and is no longer
distinguishable.[64]

7.4 Invertebrates
Arthropods have an open circulatory system, and often
some short open-ended arteries.The arthropod heart is
typically a muscular tube that runs the length of the body,
under the back and from the base of the head. Instead of
blood the circulatory uid is haemolymph which carries
the most commonly used respiratory pigment, copperbased haemocyanin as the oxygen transporter; iron-based
haemoglobin is used by only a few arthropods. The heart
contracts in ripples from the rear to the front of the ani-

16

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Basic arthropod body structure heart shown in red

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In insects, the circulatory system is not used to transport


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Additional images
The human heart viewed from the front
The human heart viewed from behind
The coronary circulation
Play media
An MRI scan showing the human heart
An anatomical specimen of the heart
Heart Illustration with circulatory system.

2011). Harrisons Principles of Internal Medicine (18


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[64] Romer, Alfred Sherwood; Parsons, Thomas S. (1977).


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10 External links
What Is the Heart? NIH
The Gross Physiology of the Cardiovascular System
(2nd Ed., 2012) Robert M. Anderson, M.D. (CCBY-NC)
Atlas of Human Cardiac Anatomy
Dissection review of the anatomy of the Human
Heart including vessels, internal and external features
Prenatal human heart development
Anatomy of the Human Heart Texas Heart Institute
Animal hearts: sh, squid

19

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11.2

Images

File:2004_Heart_Wall.jpg Source: https://upload.wikimedia.org/wikipedia/commons/3/35/2004_Heart_Wall.jpg License: CC BY 3.0


Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original artist: OpenStax
College
File:2006_Heart_Musculature.jpg Source: https://upload.wikimedia.org/wikipedia/commons/0/0e/2006_Heart_Musculature.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original
artist: OpenStax College
File:2010_Chordae_Tendinae_Papillary_Muscles.jpg Source:
https://upload.wikimedia.org/wikipedia/commons/a/a7/2010_
Chordae_Tendinae_Papillary_Muscles.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site.
http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original artist: OpenStax College
File:2011_Heart_Valves.jpg Source: https://upload.wikimedia.org/wikipedia/commons/b/b9/2011_Heart_Valves.jpg License: CC BY
3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original artist: OpenStax College

11.2

Images

21

File:2020_SA_Node_Tracing.jpg Source: https://upload.wikimedia.org/wikipedia/commons/a/a4/2020_SA_Node_Tracing.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original
artist: OpenStax College
File:2027_Phases_of_the_Cardiac_Cycle.jpg Source: https://upload.wikimedia.org/wikipedia/commons/3/38/2027_Phases_of_the_
Cardiac_Cycle.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.6/,
Jun 19, 2013. Original artist: OpenStax College
File:2028_Cardiac_Cycle_vs_Electrocardiogram.jpg Source: https://upload.wikimedia.org/wikipedia/commons/8/8d/2028_Cardiac_
Cycle_vs_Electrocardiogram.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/
col11496/1.6/, Jun 19, 2013. Original artist: OpenStax College
File:2029_Cardiac_Cycle_vs_Heart_Sounds.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/f/f3/2029_
Cardiac_Cycle_vs_Heart_Sounds.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site.
http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original artist: OpenStax College
File:2032_Automatic_Innervation.jpg Source: https://upload.wikimedia.org/wikipedia/commons/4/4c/2032_Automatic_Innervation.
jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.6/, Jun 19, 2013.
Original artist: OpenStax College
File:2037_Embryonic_Development_of_Heart.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/7/74/2037_
Embryonic_Development_of_Heart.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site.
http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original artist: OpenStax College
File:2113ab_Atherosclerosis.jpg Source: https://upload.wikimedia.org/wikipedia/commons/f/f6/2113ab_Atherosclerosis.jpg License:
CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.6/, Jun 19, 2013. Original
artist: OpenStax College
File:Apikal4D.gif Source: https://upload.wikimedia.org/wikipedia/commons/6/61/Apikal4D.gif License: CC BY-SA 3.0 Contributors:
Own work Original artist: Kjetil Lenes
File:Arthropod_body_struct_01.png Source: https://upload.wikimedia.org/wikipedia/commons/6/66/Arthropod_body_struct_01.png
License: Public domain Contributors: Transferred from en.wikipedia; transferred to Commons by User:Giancarlodessi using
CommonsHelper.
Original artist: Philcha (talk) Original uploader was Philcha at en.wikipedia
File:Commons-logo.svg Source: https://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License: ? Contributors: ? Original
artist: ?
File:ConductionsystemoftheheartwithouttheHeart-en.svg
Source:
https://upload.wikimedia.org/wikipedia/
commons/a/ae/ConductionsystemoftheheartwithouttheHeart-en.svg
License:
CC
BY-SA
3.0
Contributors:
ConductionsystemoftheheartwithouttheHeart.png & ConductionsystemoftheheartwithoutHeart.svg Original artist: Madhero88 (original les); Angelito7 (this SVG version);
File:Copy_of_Ap_Bio_2.svg Source: https://upload.wikimedia.org/wikipedia/commons/7/77/Copy_of_Ap_Bio_2.svg License: CC BYSA 4.0 Contributors: Own work Original artist: Cscanlon15
File:Coronary_arteries.svg Source: https://upload.wikimedia.org/wikipedia/commons/1/18/Coronary_arteries.svg License: CC BY-SA
3.0 Contributors:
Coronary.pdf Original artist: Coronary.pdf: Patrick J. Lynch, medical illustrator
File:Diagram_of_the_human_heart_(cropped).svg Source: https://upload.wikimedia.org/wikipedia/commons/e/e5/Diagram_of_the_
human_heart_%28cropped%29.svg License: CC-BY-SA-3.0 Contributors: Own work Original artist: ?
File:Doctors_stethoscope_1.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/e2/Doctors_stethoscope_1.jpg License:
CC BY-SA 3.0 Contributors: Own work Original artist: Stethoscopes
File:ECG_principle_slow.gif Source: https://upload.wikimedia.org/wikipedia/commons/e/e5/ECG_principle_slow.gif License: CCBY-SA-3.0 Contributors: selbst erstellt = Own work Original artist: Kalumet
File:Flow_through_the_Heart.webm
Source:
https://upload.wikimedia.org/wikipedia/commons/e/e5/Flow_through_the_
Heart.webm License: CC BY-SA 4.0 Contributors: Own work https://www.khanacademy.org/science/health-and-medicine/
human-anatomy-and-physiology/heart-introduction/v/flow-through-the-heart Original artist: Butternger8181
File:Gnome-mime-sound-openclipart.svg
Source:
https://upload.wikimedia.org/wikipedia/commons/8/87/
Gnome-mime-sound-openclipart.svg License: Public domain Contributors: Own work. Based on File:Gnome-mime-audio-openclipart.
svg, which is public domain. Original artist: User:Eubulides
File:HROgg.ogg Source: https://upload.wikimedia.org/wikipedia/commons/7/72/HROgg.ogg License: CC BY-SA 3.0 Contributors: Own
work Original artist: James Heilman, MD
File:Heart_diseccion.jpg Source: https://upload.wikimedia.org/wikipedia/commons/4/46/Heart_diseccion.jpg License: CC BY-SA 3.0
Contributors: Own work Original artist: Kenshinb
File:Latidos.gif Source: https://upload.wikimedia.org/wikipedia/commons/2/21/Latidos.gif License: Public domain Contributors: Own
work Original artist: josio
File:Leonardo_da_vinci,_Heart_and_its_Blood_Vessels.jpg
Source:
https://upload.wikimedia.org/wikipedia/commons/
c/c8/Leonardo_da_vinci%2C_Heart_and_its_Blood_Vessels.jpg License:
Public domain Contributors:
Web Gallery
of Art:
<a href='http://www.wga.hu/art/l/leonardo/10anatom/5heart.jpg' data-x-rel='nofollow'><img alt='Inkscape.svg'
src='https://upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Inkscape.svg/20px-Inkscape.svg.png'
width='20'
height='20'
srcset='https://upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Inkscape.svg/30px-Inkscape.svg.png
1.5x,
https://upload.
wikimedia.org/wikipedia/commons/thumb/6/6f/Inkscape.svg/40px-Inkscape.svg.png 2x' data-le-width='60' data-le-height='60'
/></a> Image <a href='http://www.wga.hu/html/l/leonardo/10anatom/5heart.html' data-x-rel='nofollow'><img alt='Information icon.svg'
src='https://upload.wikimedia.org/wikipedia/commons/thumb/3/35/Information_icon.svg/20px-Information_icon.svg.png' width='20'
height='20' srcset='https://upload.wikimedia.org/wikipedia/commons/thumb/3/35/Information_icon.svg/30px-Information_icon.svg.png
1.5x, https://upload.wikimedia.org/wikipedia/commons/thumb/3/35/Information_icon.svg/40px-Information_icon.svg.png 2x' data-lewidth='620' data-le-height='620' /></a> Info about artwork Original artist: Leonardo da Vinci

22

11

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

File:Real-time_MRI_-_Thorax.ogv Source: https://upload.wikimedia.org/wikipedia/commons/d/dd/Real-time_MRI_-_Thorax.ogv License: CC BY-SA 3.0 Contributors: http://www.biomednmr.mpg.de Original artist: Biomedizinische NMR Forschungs GmbH
File:Structural_organization_of_the_heart_of_the_mosquito_Anopheles_gambiae_-_image.ppat.v08.i11.g001.png
Source:
https://upload.wikimedia.org/wikipedia/commons/2/29/Structural_organization_of_the_heart_of_the_mosquito_Anopheles_gambiae_
-_image.ppat.v08.i11.g001.png License: CC BY 2.5 Contributors: King JG, Hillyer JF (2012) Infection-Induced Interaction between the
Mosquito Circulatory and Immune Systems. PLoS Pathog 8(11): e1003058. doi:10.1371/journal.ppat.1003058 Original artist: King JG,
Hillyer JF
File:Surface_anatomy_of_the_heart.png Source: https://upload.wikimedia.org/wikipedia/commons/6/6b/Surface_anatomy_of_the_
heart.png License: Public domain Contributors: See above. All used images are in public domain. Original artist: Mikael Hggstrm
File:Two_chamber_heart.svg Source: https://upload.wikimedia.org/wikipedia/commons/c/c2/Two_chamber_heart.svg License: CC BY
3.0 Contributors: Own work Original artist: Ahnode
File:Wikibooks-logo.svg Source: https://upload.wikimedia.org/wikipedia/commons/f/fa/Wikibooks-logo.svg License: CC BY-SA 3.0
Contributors: Own work Original artist: User:Bastique, User:Ramac et al.
File:Wikiquote-logo.svg Source: https://upload.wikimedia.org/wikipedia/commons/f/fa/Wikiquote-logo.svg License: Public domain
Contributors: ? Original artist: ?
File:Wikisource-logo.svg Source: https://upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg License: CC BY-SA 3.0
Contributors: Rei-artur Original artist: Nicholas Moreau
File:Wikiversity-logo-Snorky.svg Source: https://upload.wikimedia.org/wikipedia/commons/1/1b/Wikiversity-logo-en.svg License:
CC BY-SA 3.0 Contributors: Own work Original artist: Snorky
File:Wiktionary-logo-en.svg Source: https://upload.wikimedia.org/wikipedia/commons/f/f8/Wiktionary-logo-en.svg License: Public
domain Contributors: Vector version of Image:Wiktionary-logo-en.png. Original artist: Vectorized by Fvasconcellos (talk contribs),
based on original logo tossed together by Brion Vibber
File:.png Source: https://upload.wikimedia.org/wikipedia/commons/d/dd/%D0%93%D1%85%D0%B0%D0%BD%D0%B8.png
License: CC0 Contributors: Original artist:
File: -bigseal.svg Source: https://upload.wikimedia.org/wikipedia/commons/6/69/%E5%BF%83-bigseal.svg License: Public domain
Contributors: According to internationalscientic.orgs data : see L12393 image on 's page (Pinyin, English, Etymology, Cantonese,
Taiwanese informations also available). Original artist: See contributor name shown in the File history section.

11.3

Content license

Creative Commons Attribution-Share Alike 3.0

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