CHAPTER 19 : CARDIOVASCULAR

SYSTEM – HEART
FUNCTIONS OF THE Functions of Heart
1. Generating blood pressure
HEART - Contractions of the heart generate
blood pressure, which is responsible
Cardiology for moving blood through the blood
- The medical specialty concerned with vessels
diagnosing and treating heart disease 2. Routing blood
- The heart separates the pulmonary and
Pulmonary Circulation systemic circulations and ensures that
- Carries blood to the lungs, where CO2 the blood flowing to the tissues has
diffuses from the blood into the lungs and O2 adequate levels of O2
diffuses from the lungs into the blood 3. Ensuring one-way blood flow
- Returns the blood to the left side of the heart - The valves of the heart ensure a one-
way flow of blood through the heart
Systemic Circulation and blood vessels
- The left side of the heart then pumps blood 4. Regulating blood supply
delivers O2 and nutrients to all the remaining - The rate and force of heart
tissues of the body contractions change to meet the
o From those tissues, CO2 and other waste metabolic needs of the tissues, which
products are carried back to the right side vary depending on such conditions as
of the heart rest, exercise, and changes in body
position

Systemic and Pulmonary Circulation

The circulatory system consists of the pulmonary and systemic
circulations. The right side of the heart pumps blood through vessels
to the lungs and back to the left side of the heart through the
pulmonary circulation. The left side of the heart pumps blood
through vessels to the tissues of the body and back to the right side
of the heart through the systemic circulation.
 SIZE, SHAPE, AND The heart lies obliquely in the mediastinum,
with its base directed posteriorly and slightly
LOCATION OF THE HEART superiorly and its apex directed anteriorly and
slightly inferiorly. The apex is also directed to the
The adult heart is shaped like a blunt cone left, so that approximately two-thirds of the heart’s
and is approximately the size of a closed fist, with mass lies to the left of the midline of the sternum.
an average mass of 250 g in females and 300 g in The base of the heart is located deep to the sternum
males. It is larger in physically active adults and extends to the second intercostal space. The
compared with other healthy adults. The heart apex is located deep to the fifth intercostal space,
generally decreases in size after approximately age approximately 7–9 centimeters (cm) to the left of
65, especially in people who are not physically the sternum and medial to the midclavicular line, a
active. perpendicular line that extends down from the
middle of the clavicle.
Apex
- Blunt, rounded point of the heart

Base
- Larger, flat part at the opposite end of the heart

Mediastinum
- Where the heart is located
- A midline partition of the thoracic cavity that
also contains the trachea, the esophagus, the
thymus, and associated structures

It is important for health professionals to

know the location of the heart in the thoracic cavity. Location of the Heart in the Thorax
Positioning a stethoscope to hear the heart sounds (a) The heart lies in the thoracic cavity between the lungs, deep to and
slightly to the left of the sternum. The base of the heart, located deep to
and positioning electrodes to record an the sternum, extends to the second intercostal space, and the apex of the
electrocardiogram from chest leads depend on this heart is deep to the fifth intercostal space, approximately 7–9 cm to the
knowledge. left of the sternum, or where the midclavicular line intersects with the
fifth intercostal space (see inset). (b) Cross section of the thorax,
showing the position of the heart in the mediastinum and its relationship
Effective cardiopulmonary resuscitation to other structures.
(CPR) also depends on a reasonable knowledge of
the position of the heart.
 ANATOMY OF THE - Pericardial Cavity
o The space between the visceral and
HEART parietal pericardia
o Filled with a thin layer of
Pericardium serous pericardial fluid
- Pericardial sac o Helps reduce friction as the heart moves
- A double-layered, closed sac that surrounds within the pericardial sac
the heart - Even though the pericardium contains fibrous
- Consists of two layers: connective tissue, it can accommodate
1. Fibrous pericardium changes in heart size by gradually enlarging
o A tough, fibrous connective tissue o The pericardial cavity can also increase
layer that prevents overdistension in volume to hold a significant volume of
of the heart and anchors it within pericardial fluid, such as with certain
the mediastinum illnesses
o Superiorly, the fibrous
pericardium is continuous with Heart Wall
the connective tissue coverings of - Composed of three layers of tissue:
the great vessels, such as the 1. Epicardium
aorta, and inferiorly it is attached o Visceral pericardium
to the surface of the diaphragm o The superficial layer of the heart wall
2. Serous pericardium o A thin serous membrane that
o A layer of simple squamous constitutes the smooth, outer surface
epithelium of the heart
o Further subdivided into two parts: ▪ The serous pericardium is called
a. Parietal pericardium the epicardium when considered a
▪ The part lining the fibrous part of the heart and the visceral
pericardium pericardium when considered a
b. Visceral pericardium part of the pericardium
▪ Epicardium 2. Myocardium
▪ The part covering the heart o Thick, middle layer of the heart
surface o Composed of cardia muscle cells
o Responsible for the heart’s ability to
contract
3. Endocardium
o Deep to the myocardium
o Consists of simple squamous
epithelium over a layer of connective
tissue
o Form the smooth, inner surface of the
heart chambers, which allows blood to
move easily through the heart
o Covers the surfaces of the heart valves

Heart in the Pericardium Heart Wall

The heart is surrounded by the pericardium, which consists of an outer Part of the wall of
fibrous pericardium and an inner serous pericardium. The serous the heart has been
pericardium has two parts: The parietal pericardium lines the fibrous removed, enlarged,
pericardium, and the visceral pericardium (epicardium) covers the and rotated, so that
surface of the heart. The pericardial cavity, between the parietal and its inner surface is
visceral pericardia, is filled with a small amount of pericardial fluid. visible. The
enlarged section
- The parietal and visceral portions of the illustrates the
serous pericardium are continuous with each epicardium
(visceral
other where the great vessels enter or leave pericardium),
the heart myocardium, and
endocardium.
 Ridges formed by the myocardium can be
seen on the internal surfaces of the heart chambers.
Though the interior surfaces of the atria are mainly
flat, the interior of both auricles and a part of the
right atrial wall contain muscular ridges
called pectinate muscles. The pectinate muscles
of the right atrium are separated from the larger,
smooth portions of the atrial wall by a ridge called
the crista terminalis (terminal crest). The interior
walls of the ventricles contain larger, muscular
ridges and colums called trabeculae carneae.
These ridges help with forceful ejection of blood (b) Anterior View
from the ventricles.

External Anatomy and Coronary Circulation

- The heart consists of four chambers: (c) Posterior View
o Two Atria Surface View of the Heart
▪ Thin-walled (a) The two atria (right and left) are located superiorly, and the two ventricles (right
and left) are located inferiorly. The superior and inferior venae cavae enter the right
▪ Forms the superior and posterior atrium. The pulmonary veins enter the left atrium. The pulmonary trunk exits the
parts of the heart right ventricle, and the aorta exits the left ventricle.
(b) Photograph of the anterior surface of the heart. (c) The two atria (right and left)
o Two Ventricles are located superiorly, and the two ventricles (right and left) are located inferiorly.
▪ Thick-walled The superior and inferior venae cavae enter the right atrium, and the four pulmonary
veins enter the left atrium. The pulmonary trunk divides, forming the left and right
▪ Forms the anterior and inferior pulmonary arteries.
portions
- Auricles - Blood enters the atria of the heart through
o Flaplike extensions of the atria that can several large veins
be seen anteriorly between each atrium o The Superior vena cava and the Inferior
and ventricle vena cava carry blood from the body to
the right atrium
o In addition, the smaller coronary sinus
carries blood from the walls of the heart to
the right atrium
o Four Pulmonary veins carry blood from
the lungs to the left atrium
- Blood leaves the ventricles of the heart
through two arteries:
o Pulmonary Trunk
▪ Carries blood from the right ventricle
to the lungs
o Aorta
▪ Carries blood from the left ventricle
to the body
o Because of their large size, the
(a) Anterior View pulmonary trunk and aorta are often
called the great arteries
- The coronary circulation consists of blood
vessels that carry blood to and from the
tissues of the heart wall
o The major vessels of the coronary
circulation lie in several grooves, or sulci,
on the surface of the heart
o Coronary Sulcus
▪ Large groove
▪ Runs obliquely around the heart,
separating the atria from the
ventricles
o Two more grooves extend inferiorly from
the coronary sulcus, indicating the
division between the right and left
(b)
ventricles:
▪ Anterior interventricular sulcus Coronary Circulation
(a) Arteries supplying blood to the heart. The arteries of the anterior
• On the anterior surface of the surface are seen directly and are darker in color; the arteries of the
heart, extending from the posterior surface are seen through the heart and are lighter in color.
(b) Veins draining blood from the heart. The veins of the anterior
coronary sulcus toward the surface are seen directly and are darker in color; the veins of the
apex of the heart posterior surface are seen through the heart and are lighter in color.

▪ Posterior interventricular sulcus

- The Left coronary artery has three major
• On the posterior surface of the branches:
heart, extending from the 1. Anterior interventricular artery
coronary sulcus toward the o Left anterior descending artery
apex of the heart o First major branch
o In a healthy, intact heart, the grooves are o Extends inferiorly in the anterior
covered by adipose tissue, and only after interventricular sulcus and supplies
this tissue is removed can they be seen blood to most of the anterior part of
- The major arteries supplying blood to the the heart
tissue of the heart lie within the coronary 2. Left marginal artery
sulcus and interventricular grooves on the o Second major branch
surface of the heart o Supplies blood to the lateral wall of
o The Right and Left coronary the left ventricle
arteries exit the aorta just above the 3. Circumflex artery
point where the aorta leaves the heart o Third major branch
▪ These vessels lie within the coronary o Extends around to the posterior side of
sulcus the heart in the coronary sulcus
▪ The right coronary artery is usually o Supply blood to much of the posterior
smaller in diameter than the left one, wall of the heart
and it does not carry as much blood - The Right coronary artery has two major
as the left coronary artery branches:
o The right coronary artery lies within the
coronary sulcus and extends from the
aorta around to the posterior part of the
heart
1. Right marginal artery
▪ Larger branch
▪ Supply blood to the lateral wall of
the right ventricle
2. Posterior interventricular artery
▪ Second branch
▪ Lies in the posterior
interventricular sulcus
(a) ▪ Supplies blood to the posterior
and inferior part of the heart
- Most of the myocardium receives blood from ▪ A number of smaller veins empty into
more than one arterial branch the cardiac veins, into the coronary
o In addition, the coronary circulation sinus, or directly into the right atrium
includes many Anastamoses, or direct - Blood flow through the coronary blood
connections between arteries circulation is not continuous
▪ These anastamoses may form either o When the cardiac muscle contracts, blood
between branches of a given artery vessels in the wall of the heart are
or between branches of different compressed so blood does not readily
arteries flow through them
▪ As a result of these many o When the cardiac muscle relaxes, the
connections among the coronary blood vessels are not compressed, and
arteries, if one artery becomes blood flow through the coronary blood
blocked, the areas primarily supplied vessels resumes.
by that artery may still receive some - As blood flows through tissues, O2 is released
blood through other arterial branches from the blood and moves into the tissues
and anastamoses o The amount of O2 released varies from
▪ The density of blood vessels one tissue to another, even in the case of
supplying blood to the myocardium the different muscle tissue types
increases with aerobic exercise, as do o Cardiac muscle requires blood to flow
the number and extent of the through the coronary arteries at a higher
anastamoses rate than its resting level in order to
provide an adequate O2 supply during
exercise

Heart Chambers and Valves

1. Right and Left Atria
- Right Atrium
o Has three major openings:
▪ An opening from the
superior vena cava
▪ An opening from the
- The coronary circulation also includes veins inferior vena cava
that carry the blood from the heart walls to ▪ An opening from the
the right atrium coronary sinus
o Two major veins draining the blood from o The openings from the superior
the heart wall tissue: vena cava and the inferior vena
1. Great cardiac vein cava receive blood from the
▪ Drains blood from the left side
body, and the opening of the
of the heart coronary sinus receives blood
2. Small cardiac vein
from the heart itself
▪ Drains the right margin of the - Left Atrium
heart o Has four relatively uniform
openings from the four
pulmonary veins that receive
blood from the lungs

o These veins converge toward the

posterior part of the coronary sulcus and
empty into a large venous cavity called
the Coronary sinus, which in turn
empties into the right atrium
 - Interatrial septum - Bicuspid valve
o A wall of tissue that separates o mitral valve
the right and left atria from each o The atrioventricular valve
other between the left atrium and the
- Fossa ovalis left ventricle
o A slight, oval depression on the o It has two cusps
right side of the interatrial
septum marking the former
location of the foramen ovale
- Foramen ovale
o An opening between the right
and left atria in the embryonic
and fetal heart
o In the fetal heart, this opening
allows blood to flow from the
right to the left atrium and (a) Anterior View
bypass the pulmonary
circulation
2. Right and Left Ventricles
- The atria open into the ventricles
through Atrioventricular canals
- Each ventricle has one large,
superiorly placed outflow route near
the midline of the heart
- Blood flows from the Right
ventricle into the pulmonary trunk
- Blood flows from the Left (b) Superior View
ventricle into the aorta
Heart Valves
- Interventricular septum (a) Tricuspid valve, chordae tendineae, and papillary muscles. (b)
o The one that separates the two Heart valves. Note the three cusps of each semilunar valve meeting
ventricles from each other to prevent the backflow of blood.
o Has a thick, muscular part
toward the apex and a thin,
- Papillary muscles
membranous part toward the
o Cone-shaped, muscular pillars
atria
that each ventricle contains
- The wall of the left ventricle is much
o These muscles are attached to
thicker than the wall of the right
the cusps of the atrioventricular
ventricle
valves by thin, strong connective
o The thicker wall of the left
tissue strings called chordae
ventricle allows for stronger
tendineae
contractions to pump blood
o It contracts when the ventricles
through the systemic circulation
contract and prevent the valves
3. Atrioventricular Valves
from opening into the atria by
- Is in each atrioventricular canal and
pulling on the chordae tendineae
is composed of cusps, or flaps
attached to the valve cusps
- Ensure blood flows from the atria
- The atrioventricular canal is closed
into the ventricles, preventing blood
as the valve cusps meet
from flowing back into the atria
- Tricuspid valve
o The atrioventricular valve
between the right atrium and the
right ventricle
o Consists of three cusps
 4. Semilunar Valves
- Positioned between each ventricle and
its associated great artery
- Are identified by the great artery in
which each is located and include the
Aortic semilunar valve and Pulmonary
semilunar valve
o Each valve consists of three
pocketlike, semilunar cusps, the
free inner borders of which meet
in the center of the artery to block
blood flow
- Contraction of the ventricles pushes
blood against the semilunar valves,
forcing them to open
o Blood can then enter the great
arteries
o However, when blood flows back
from the aorta or pulmonary trunk
toward the ventricles, it enters the
pockets of the cusps, causing the
cusps to meet in the center of the
aorta or pulmonary trunk
▪ This effectively closes the
semilunar valves and
prevents blood from flowing
back into the ventricles

Function of the Heart Valves

(a) Valve positions when blood is flowing into the left
ventricle. (b) Valve positions when blood is flowing out of
the left ventricle. Numbered steps show the functions of the
bicuspid and aortic semilunar valves. The tricuspid and
pulmonary semilunar valves (not shown) open and close in
a similar pattern.
 ROUTE OF BLOOD FLOW o Blood also pushes against the pulmonary
semilunar valve, forcing it open
THROUGH THE HEART o Blood then flows into the pulmonary
trunk
Blood Flow
- Blood flows through both sides The pulmonary trunk branches to form
simultaneously the Pulmonary arteries, which carry blood to the
- Both atria contract at about the same time and lungs, where CO2 is released and O2 is picked
both ventricles contract at about the same up. Blood returning from the lungs enters the left
time, therefore blood is moving through both atrium through the four pulmonary veins. Most of
the pulmonary and the systemic circulations the blood passes from the left atrium into the
with each heartbeat relaxed left ventricle. Contraction of the left atrium
completes left ventricular filling.

Contraction of the left ventricle pushes blood

against the bicuspid valve, closing it and
preventing blood from moving back into the left
atrium. Blood is also pushed against the aortic
semilunar valve, opening it and allowing blood to
enter the aorta. Blood flowing through the aorta is
distributed to all parts of the body, except to the
parts of the lungs supplied by the pulmonary blood
vessels.
Recall that in the embryonic and fetal heart the
foramen ovale allows for blood to flow between the
two atria. This hole closes at birth, separating the
right and left sides of the heart.

Blood Flow Through the Heart

(a) Frontal section of the heart revealing the four chambers and the
direction of blood flow (purple numbers). (b) Diagram listing, in order,
the structures through which blood flows in the systemic, pulmonary,
and coronary circulations. The heart valves are indicated
by circles; deoxygenated blood appears blue, and oxygenated blood
appears red.

- Blood enters the relaxed right atrium from the

systemic circulation, which returns blood
from all the tissues of the body
o Most of the blood in the right atrium then
passes into the relaxed right ventricle
o The right atrium then contracts, pushing
most of the remaining blood in the atrium
into the right ventricle to complete right
ventricular filling
- Contraction of the right ventricle pushes
blood against the tricuspid valve, forcing it
closed
o Closing of the tricuspid valve prevents
blood from moving back into the right
atrium
 HISTOLOGY
Heart Skeleton
- Consists of a plate of fibrous connective
tissue between the atria and the ventricles
o This connective tissue plate forms fibrous
rings around the atrioventricular and
semilunar valves and provides solid
support for them, reinforcing the valve
openings
o The fibrous connective tissue plate also
serves as electrical insulation between the
atria and the ventricles and provides a rigid
site for attachment of the cardiac muscles Histology of the Heart
(a) Cardiac muscle cells are branching cells with centrally located nuclei.
The cells are joined to one another by intercalated disks. Gap junctions
in the intercalated disks allow action potentials to pass from one cardiac
muscle cell to the next. (b) A light micrograph of cardiac muscle tissue.
The cardiac muscle cells appear striated because of the arrangement of
the individual myofilaments. (c) As in skeletal muscle, sarcomeres join
end-to-end to form myofibrils, and mitochondria provide ATP for
contraction. Sarcoplasmic reticulum and T tubules are visible but are not
as numerous as they are in skeletal muscle.

- Cardiac muscle cell contraction is very

similar to that of skeletal muscle; however,
the onset of contraction is longer and
prolonged in cardiac muscle
o These differences in contraction are
Heart Skeleton partially due to differences in cell
The skeleton of the heart consists of fibrous connective anatomy
tissue rings, which surround the heart valves and separate o Cardiac muscle has a
the atria from the ventricles. Cardiac muscle attaches to
the fibrous connective tissue. The muscle fibers are smooth Sarcoplasmic reticulum
arranged so that contraction of the ventricles produces a ▪ Stores Ca2+, similar to skeletal
wringing motion and the distance between the apex and
the base of the heart shortens.
muscle
▪ It is not as regularly arranged as in
skeletal muscle fibers, and there are
Cardiac Muscle no dilated cisternae, as in skeletal
- Cardiac muscle cells muscle
o Elongated, branching cells that have one,
or occasionally two, centrally located
nuclei
o Contain actin and myosin myofilaments
organized to form sarcomeres, which join
end-to-end to form myofibrils
▪ The actin and myosin myofilaments
are responsible for cardiac muscle
contraction, and their organization
gives cardiac muscle a striated
(banded) appearance
▪ Striations are less regularly arranged
- Cardiac muscles have transverse tubules that
and less numerous than in skeletal
are in close association with the sarcoplasmic
muscle
reticulum
o However, the T tubules in cardiac muscle
are larger in diameter than in skeletal
muscle, and extensions of T tubules are
not as closely associated with the
 sarcoplasmic reticulum as in skeletal to-end; however, intercalated disks
muscle can also connect cells laterally
o The T tubules of cardiac muscle are o At intercalated disks, the plasma
found near the Z disks of the sarcomeres, membranes are folded, and the adjacent
instead of where the actin and myosin cells fit together, thus greatly increasing
overlap, as in skeletal muscle contact between them
▪ Given these structural differences, o Specialized plasma membrane structures
depolarizations of the cardiac muscle at the intercalated disks increase physical
plasma membrane are not carried and electrical connections between cells
from the surface of the cell to the ▪ These plasma membrane structures
sarcoplasmic reticulum as efficiently include desmosomes and gap
as they are in skeletal muscle, and junctions
Ca2+ must diffuse a greater distance o Desmosomes hold the cells together,
from the sarcoplasmic reticulum to and gap junctions allow cytoplasm to
the actin myofilaments flow freely between cells, resulting in
o Another important difference between areas of low electrical resistance between
cardiac muscle and skeletal muscle is the the cells
sources for Ca2+ necessary for ▪ This enables action potentials to pass
contraction easily from one cell to the next
▪ In skeletal muscle, adequate Ca2+ for o Electrically, the cardiac muscle cells
contraction is stored in the behave as a single unit, and the heart’s
sarcoplasmic reticulum, but cardiac highly coordinated contractions depend
muscle requires some Ca2+ from the on this functional characteristic
extracellular fluid and from the T
tubules
- Cardiac muscle is specialized to meet the high
energy requirements needed for proper
myocardial function
o ATP provides the energy for cardiac
muscle contraction
o ATP production depends on
O2 availability
▪ Because cardiac muscle must
continue to contract and relax in a Conducting System
relatively steady rhythm to maintain - Relays action potentials through the heart
life, it cannot develop a large oxygen - Consists of modified cardiac muscle cells that
deficit, which is often seen in form two nodes (knots or lumps) and a
skeletal muscle conducting bundle
o Cardiac muscle cells are rich in o The two nodes are contained within the
mitochondria, which perform oxidative walls of the right atrium and are named
metabolism at a rate rapid enough to according to their position in the atrium
sustain normal myocardial energy - The two nodes:
requirements 1. Sinoatrial (SA) node
▪ Also, the myocardium has an o The medial to the opening of the
extensive capillary network that superior vena cava
provides an adequate O2 supply to 2. Atrioventricular (AV) node
the cardiac muscle cells. o The medial to the right atrioventricular
- Cardiac muscle cells are organized in spiral valve
bundles or sheets o Gives rise to a conducting bundle of
o Cardiac cells are bound to adjacent cells the heart, the Atrioventricular (AV)
by specialized cell-to-cell contacts bundle (bundle of His)
called Intercakated disks ▪ This bundle passes through a small
▪ Intercalated disks are located at the opening in the fibrous skeleton to
ends of cells, connecting them end- reach the interventricular septum,
where it divides to form the right
 and left bundle branches, which - Unlike skeletal muscle cells that require
extend beneath the endocardium on neural stimulation for contraction, cardiac
each side of the interventricular muscle cells have the intrinsic capacity to
septum to the apex of both the right spontaneously generate action potentials for
and the left ventricles contraction
o Because cells of the SA node
spontaneously generate action potentials
at a greater frequency than other cardiac
muscle cells, these cells are called
the pacemaker of the heart
o The SA node is made up of specialized,
small-diameter cardiac muscle cells that
merge with the other cardiac muscle cells
of the right atrium
▪ Once action potentials are produced,
they spread from the SA node to
adjacent cardiac muscle cells of the
atrium
Conducting System of the Heart o Preferential pathways conduct action
The conduction system of the heart is composed of specialized cardiac potentials from the SA node to the AV
muscle cells that produce spontaneous action potentials. The organization node at a greater velocity than they are
of the conduction system ensures the proper pattern of contractions of the
atria and ventricle, maintaining normal blood flow. transmitted in the remainder of the atrial
muscle cells
- Purkinje fibers ▪ However, such pathways cannot be
o The inferior terminal branches of the distinguished structurally from the
bundles remainder of the atrium
o Are large-diameter cardiac muscle fibers o It is the activity of the SA node that
o They have fewer myofibrils than most causes the heart to contract
cardiac muscle cells and do not contract spontaneously and rhythmically
as forcefully - When the heart beats under resting
o Intercalated disks are well developed conditions, approximately 0.04 second is
between the Purkinje fibers and contain required for action potentials to travel from
numerous gap junctions the SA node to the AV node
▪ As a result of these structural o Action potentials are propagated slowly
modifications, action potentials through the AV node, compared with the
travel along the Purkinje fibers much remainder of the conducting system
more rapidly than through other o The slow rate of action potential
cardiac muscle tissue conduction in the AV node is due, in part,
to the smaller-diameter muscle cells and
fewer gap junctions in their intercalated
disks
▪ Just like other specialized conducting
cells in the heart, they have fewer
myofibrils than most cardiac muscle
cells
• As a consequence, a delay of
0.11 second occurs from the
time action potentials reach the
AV node until they pass to the
AV bundle
• The delay of action potentials at
the AV node allows for
completion of the atrial
contraction before ventricular
contraction begins
 o After action potentials pass from the AV
node to the highly specialized conducting
bundles, the velocity of conduction
increases dramatically
▪ The action potentials pass through
the left and right bundle branches
and through the individual Purkinje
fibers that penetrate the myocardium
of the ventricles
- Because of the arrangement of the conducting
system in the ventricles, the first part of the
ventricular myocardium that is stimulated is
the inner wall of the ventricles near the apex
o Thus, ventricular contraction begins at
the apex and progresses throughout the
ventricles toward the base of the heart
o The spiral arrangement of muscle layers
in the wall of the heart results in a
wringing action
o During the process, the distance between
the apex and the base of the heart
decreases and blood is forced upward
from the apex toward the great vessels at
the base of the heart
 ELECTRICAL Comparison of Action Potentials in Skeletal and Cardiac
Muscle
PROPERTIES (a) An action potential in skeletal muscle consists of
depolarization and repolarization phases. (b) An action potential
in cardiac muscle consists of depolarization, early repolarization,
plateau, and final repolarization phases. Cardiac muscle does not
Cardiac muscle cells—like other repolarize as rapidly as skeletal muscle (indicated by the break in
electrically excitable cells, such as neurons and the curve) because of the plateau phase.
skeletal muscle fibers—have a resting membrane
potential, the membrane potential when the cell is - The longer action potentials in cardiac muscle
relaxed. The resting membrane potential depends can be divided into four phases, each
on a low permeability of the plasma membrane to associated with specific changes in ion
Na+ and Ca2+ and a higher permeability to K+. movement across the membrane
When neurons, skeletal muscle fibers, and cardiac 1. A rapid depolarization phase during
muscle cells are depolarized to their threshold which the membrane potential quickly
level, action potentials result becomes more positive
2. A rapid but partial early repolarization
Action Potentials phase
- Cardiac muscle exhibit depolarization 3. A prolonged period of slow repolarization,
followed by repolarization of the the plateau phase
resting membrane potential 4. A more rapid final repolarization
- Alterations in membrane channels are phase, during which the membrane
responsible for the changes in the potential returns to its resting level
permeability of the plasma membrane that - Rapid depolarization
produce the action potentials o The result of changes in membrane
- Action potentials in cardiac muscle last longer permeability to Na+, K+, and Ca2+
than those in skeletal muscle, and the o Membrane permeability to Na+ is the
membrane channels differ somewhat from primary determinant of this phase
those in skeletal muscle o Membrane channels, called voltage-gated
o In contrast to action potentials in skeletal Na+ channels, open, bringing about the
muscle, which take less than 2 depolarization phase of the action
milliseconds (ms) to complete, action potential
potentials in cardiac muscle take o As the voltage-gated Na+channels open,
approximately 200–500 ms to complete Na+ diffuses into the cell, causing rapid
depolarization until the cell is
depolarized to approximately +20
millivolts (mV)
- The voltage change occurring during
depolarization affects other ion channels in
the plasma membrane
o Several types of voltage-gated
K+channels exist, each of which opens
and closes at different membrane
potentials, causing changes in membrane
permeability to K+
o Depolarization causes these voltage-
gated K+ channels to close, thereby
decreasing membrane permeability to K+
▪ It also causes voltage-gated
Ca2+ channels to begin to open
• These changes contribute to
depolarization
o Compared with Na+ channels, the
Ca2+ channels open and close slowly
- Repolarization
o The result of changes in membrane
permeability to Na+, K+, and Ca2+
 o Early repolarization occurs when the muscle cells stimulates the release of
voltage-gated Na+ channels and some Ca2+ from the sarcoplasmic reticulum, a
voltage-gated Ca2+ channels close, and a process called calcium-induced calcium
small number of voltage-gated release (CICR)
K+ channels open ▪ When an action potential occurs in a
o Sodium ion movement into the cell cardiac muscle cell, Ca2+ enters the
slows, and some K+ moves out of the cell cell and binds to receptors in the
▪ At this point, repolarization begins, membranes of the sarcoplasmic
but in cardiac muscle early reticulum, resulting in the opening of
repolarization is slow due to the Ca2+ channels on the membrane of
influx of Ca2+, resulting in a plateau the sarcoplasmic reticulum
phase ▪ Calcium ions then move out of the
o The plateau phase occurs as voltage- sarcoplasmic reticulum and activate
gated Ca2+ channels remain open, and the interaction between actin and
Ca2+ and some Na+ move into the cell myosin to produce contraction of the
through the voltage-gated Ca2+ channels cardiac muscle cells
▪ The influx of these ions counteracts o Skeletal muscle contraction does not
the potential change produced by the depend on this mechanism and relies only
movement of K+ out of the cell on intracellular Ca2+ for contraction
▪ The plateau phase ends, and final
repolarization begins as the voltage- Autorhythmicity of Cardiac Muscle
gated Ca2+ channels close and many - The heart is Autorhythmic
more voltage-gated K+ channels open o It stimulates itself to contract at regular
• Thus, Ca2+ and Na+ stop diffusing intervals
into the cell, and the tendency for - In the SA node, pacemaker cells generate
K+ to diffuse out of the cell action potentials spontaneously and at regular
increases intervals
o These permeability changes cause the o These action potentials spread through
membrane potential to return to its the conducting system of the heart to
resting level other cardiac muscle cells, causing
- Action potential propagation in cardiac voltage-gated Na+ channels to open
muscle differs from that in skeletal muscle ▪ As a result, action potentials are
o First, action potentials in cardiac muscle produced and the cardiac muscle
are conducted from cell to cell through cells contract
the gap junctions of the intercalated - Depolarization of pacemaker cells is
disks, whereas action potentials in dependent on Na+, K+, and Ca2+; however, the
skeletal muscle fibers are conducted ways that these ions affect the membrane
along the length of a single muscle fiber potential are very different
(cell), but not from fiber to fiber o When a spontaneously developing local
o Second, action potential propagation is potential, called the pacemaker
slower in cardiac muscle than in skeletal potential, reaches threshold, then action
muscle because cardiac muscle cells are potentials are generated in the SA node
smaller in diameter and much shorter ▪ Changes in ion movement into and
than skeletal muscle fibers out of the pacemaker cells cause the
o Although the gap junctions allow the pacemaker potential
transfer of action potentials between o Sodium ions cause depolarization by
cardiac muscle cells, they slow the rate of moving into the cells through specialized
action potential conduction between the nongated Na+ channels
cardiac muscle cells o A decreasing permeability to K+ also
- Another interesting characteristic of cardiac causes depolarization as less K+moves
muscle contraction is the need for out of the cells
extracellular Ca2+ for contraction to occur ▪ The decreasing K+ permeability
o The movement of Ca2+ through the occurs due to the voltage changes at
plasma membrane, including the the end of the previous action
membranes of the T tubules, into cardiac potential
 ▪ As a result of the depolarization, ➢ For example, if the SA node
voltage-gated Ca2+ channels open, does not function properly, the
and the movement of Ca2+ into the part of the heart that can
pacemaker cells causes further produce action potentials at the
depolarization next highest frequency is the
o When the pacemaker potential reaches AV node, which produces a
threshold, many voltage-gated heart rate of 40–60 bpm
Ca2+ channels open • Another cause of an ectopic focus
▪ In pacemaker cells, the movement of is blockage of the conducting
Ca2+ into the cells is primarily pathways between the SA node
responsible for the depolarization and other parts of the heart
phase of the action potential ➢ For example, if
▪ This is different from other cardiac action potentials do not pass
muscle cells, where the movement of through the AV node, an
Na+ into the cells is primarily ectopic focus can develop in an
responsible for depolarization AV bundle, resulting in a heart
o Repolarization occurs, as in other cardiac rate of only 30 bpm
muscle cells, when the voltage-gated • Can also appear when the rate of
Ca2+ channels close and the voltage-gated action potential generation in
K+ channels open cardiac muscle cells outside the
o After the resting membrane potential is SA node becomes enhanced
reestablished, production of another ➢ For example, when cells are
pacemaker potential starts the generation injured, their plasma
of the next action potential membranes become more
permeable, resulting in
depolarization. Inflammation
or lack of adequate blood flow
to cardiac muscle tissue can
injure cardiac muscle cells
➢ These injured cells can be the
source of ectopic action
potentials
• Changes in cardiac muscle cells’
Pacemaker Potential membrane potentials or
The production of action potentials by the pacemaker cells of the SA permeability can produce ectopic
node is responsible for the autorhythmicity of the heart.
foci
- Although most cardiac muscle cells respond ➢ Alterations in blood levels of
to action potentials produced by the SA node, K+and Ca2+ can change the
some cardiac muscle cells in the conducting cardiac muscle membrane
system can also generate spontaneous action potential, and certain drugs,
potentials such as those that mimic the
o Normally, the SA node controls the effect of epinephrine on the
rhythm of the heart because its heart, can alter cardiac muscle
pacemaker cells generate action membrane permeability
potentials at a faster rate than other
potential pacemaker cells Refractory Periods of Cardiac Muscle
o The SA node produces a heart rate of 70– - Cardiac muscle, like skeletal muscle,
80 beats per minute (bpm) has Refractory periods associated with its
o In some conditions, another area of the action potentials
conducting system may generate a - The refractory period can be subdivided into
heartbeat the absolute refractory period and the relative
▪ An Ectopic Focus refractory period
• Any part of the heart other than 1. Absolute refractory period
the SA node that generates a o The cardiac muscle cell is completely
heartbeat insensitive to further stimulation
 2. Relative refractory period important electrical changes of the
o The cell is sensitive to stimulation, but a myocardium of the heart
greater stimulation than normal is 1. P wave
required to cause an action potential o The result of action potentials that
- Because the plateau phase of the action cause depolarization of the atrial
potential in cardiac muscle delays myocardium, signals the inset of atrial
repolarization to the resting membrane contraction
potential, the refractory period is prolonged 2. QRS complex
- The long refractory period ensures that o Composed of three individual
contraction and most of relaxation are waves—the Q, R, and S waves—
complete before another action potential can results from ventricular depolarization
be initiated and signals the onset of ventricular
o This prevents tetanic contractions in contraction
cardiac muscle and is responsible for 3. T wave
rhythmic contractions o Represents repolarization of the
ventricles and precedes ventricular
Electrocardiogram relaxation
- Action potentials conducted through the o A wave representing repolarization of
myocardium during the cardiac cycle produce the atria cannot be seen because it
electrical currents that can be measured at the occurs during the QRS complex
body surface - In addition to the different waves that can be
- Electrodes placed on the body surface and detected on an ECG, time intervals can also
attached to an appropriate recording device be determined
can detect small voltage changes resulting o The time between the beginning of the P
from action potentials in the cardiac muscle wave and the beginning of the QRS
o The electrodes do not detect individual complex is the PQ interval, commonly
action potentials; rather, they detect a called the PR interval because the Q
summation of all the action potentials wave is often very small
transmitted by the cardiac muscle cells o During the PR interval, which lasts
through the heart at a given time approximately 0.16 second, the atria
- Electrocardiogram (ECG or EKG) contract and begin to relax
o The summated record of the cardiac action ▪ The ventricles begin to depolarize at
potentials the end of the PR interval
o A record of the electrical activity of the o The QT interval extends from the
heart beginning of the QRS complex to the end
o Not a direct measurement of mechanical of the T wave, lasting approximately 0.36
events in the heart, neither the force of second
contraction nor blood pressure can be ▪ The QT interval represents the
determined from it approximate length of time required
o Each deflection in the ECG record for the ventricles to contract and
indicates an electrical event within the begin to relax
heart that is correlated with a subsequent - Elongation of the PR interval can result
mechanical event from three events:
o ECG analysis can reveal abnormal 1. A delay in action potential conduction
conduction pathways, hypertrophy or through the atrial muscle because of
atrophy of portions of the heart, and the damage, such as that caused by Ischemia,
approximate location of damaged cardiac which is obstruction of the blood supply
muscle to the walls of the heart
- Electrocardiography 2. A delay in action potential conduction
o Extremely valuable in diagnosing a through atrial muscle because of a dilated
number of abnormal cardiac rhythms and atrium
other abnormalities, particularly because it 3. A delay in action potential conduction
is painless, easy to record, and noninvasive through the AV node and bundle because
- The normal ECG consists of a P wave, a QRS of ischemia, compression, or necrosis of
complex, and a T wave, each representing the AV node or bundle
 o These conditions result in slow
conduction of action potentials through
the bundle branches
o An unusually long QT interval reflects
the abnormal conduction of action
potentials through the ventricles, which
can result from myocardial infarctions or
from an abnormally enlarged left or right
ventricle
- Altered forms of the electrocardiogram due to
cardiac abnormalities include complete heart
block, premature ventricular contraction,
bundle branch block, atrial fibrillation, and
ventricular fibrillation

Alterations in an Electrocardiogram
Abnormal conduction of action potentials through the
conducting system and the myocardium leads to
alterations of patterns of an ECG
 CARDIAC CYCLE o It can be as short as 0.25–0.3 second in a
newborn or as long as 1 or more seconds
in a well-trained athlete
The right and left halves of the heart can be
o The normal cardiac cycle of 0.7–0.8
viewed as two separate pumps that work together. second depends on the capability of
Each pump consists of a “primer pump” (the cardiac muscle to contract and on the
atrium) and a “power pump” (the ventricle). Both
functional integrity of the conducting
atrial primer pumps complete the filling of the system
ventricles with blood, and both ventricular power
- The cardiac cycle involves a predictable
pumps produce the major force that causes blood to pattern of contraction and relaxation of the
flow through the pulmonary and systemic arteries. heart chambers
o Systole
Cardiac Cycle
▪ Means to contract
- Refers to the repetitive pumping process that
o Diastole
begins with the onset of cardiac muscle ▪ Means to dilate
contraction and ends with the beginning of o Atrial Systole
the next contraction
▪ Contraction of the atrial myocardium
- Blood moves from an area of higher pressure o Atrial Diastole
to an area of lower pressure
▪ Relaxation of the atrial myocardium.
o Pressure changes produced within the o Ventricular Systole
heart chambers as a result of cardiac ▪ Contraction of the ventricular
muscle contraction and relaxation move myocardium
blood along the previously described o Ventricular Diastole
routes of the pulmonary and systemic ▪ Relaxation of the ventricular
circulations myocardium
▪ When the
terms systole and diastole are used
alone, they refer to ventricular
systole and diastole
- At the beginning of the cardiac cycle, the atria
and ventricles are relaxed, the AV valves are
open, and the semilunar valves are closed
o During the cardiac cycle, changes in
chamber pressure and the opening and
closing of the heart valves determine the
direction of blood movement
o It is important to focus on these pressure
changes and heart valve movements
- At rest, most of the blood movement into the
chambers is a passive process resulting from
the greater blood pressure in the veins than in
the heart chambers
o As the blood moves into the atria, much
of it flows into the ventricles for two
reasons:
▪ The AV valves are open
▪ The atrial pressure is slightly greater
than ventricular pressure
o Passive ventricular filling
Cardiac Cycle ▪ The time period when blood is
The cardiac cycle is a repeating series of contraction and relaxation that passively moving into the ventricles
moves blood through the heart (AV = atrioventricular).
- A detailed description of the cardiac cycle
- The duration of the cardiac cycle varies describing the stimulation of the heart
considerably among humans and during an chambers, changes in pressure, and opening
individual’s lifetime and closing of the heart valves:
1. Active ventricular filling o As ventricular diastole begins, the
o STEP 1 ventricles relax, and ventricular
o The SA node generates an action pressures decrease below the pressures
potential that stimulates atrial in the pulmonary trunk and aorta
contraction o Consequently, blood begins to flow
o This P wave of an ECG represents this back toward the ventricles, causing the
electrical activity semilunar valves to close
o Atrial contraction begins the cardiac o With closure of the semilunar valves, all
cycle the heart valves are closed, and no
▪ As the atria contract, they carry out blood flows into the relaxing ventricles
the primer pump function by 5. Atrial diastole began during ventricular
forcing more blood into the systole, and as the atria relaxed, blood
ventricles flowed into them from the veins
2. Period of isovolumetric contraction o STEP 5
o STEP 2 o As the ventricles continue to relax,
o The action potential passes to the AV ventricular pressures drop below atrial
node, down the AV bundle, bundle pressures, and the AV valves open
branches, and Purkinje fibers, o Passive ventricular filling begins again
stimulating ventricular systole - Once the ventricles have fully relaxed, the
▪ This electrical activity is state of the heart is the same as when the
represented as the QRS complex of cardiac cycle began, all chambers are relaxed,
an ECG the AV valves are open, and the semilunar
o As the ventricles contract, ventricular valves are closed
pressures increase, causing blood to o With the next stimulus from the SA node,
flow toward the atria and close the AV another cardiac cycle will begin
valves
▪ The semilunar valves are closed at Events Occurring During the Cardiac Cycle
this point as well - Main events of the cardiac cycle should be
▪ Ventricular contraction continues examined from top to bottom:
and ventricular pressures rise; o Panel 1:
however, because all the valves are ▪ An ECG indicates the electrical events
closed, no blood flows from the that cause contraction and relaxation of
ventricles at this time the atria and ventricles
o Brief interval o Panel 2:
▪ The pressure graph shows the pressure
▪ Because the volume of blood in the
changes within the left atrium, left
ventricles does not change, even ventricle, and aorta resulting from atrial
though the ventricles are and ventricular contraction and
contracting relaxation
3. Period of ejection ▪ Although pressure changes in the right
o STEP 3 side of the heart are not shown, they are
o Ventricular contraction continues, and similar to those in the left side, only
ventricular pressure builds until it lower
overcomes the pressures in the o Panel 3:
pulmonary trunk and aorta ▪ The volume graph presents the changes
▪ As a result, the semilunar valves in left ventricular volume as blood flows
are pushed open, and blood flows into and out of the left ventricle as a
result of the pressure changes
from the ventricles into those
o Panel 4:
arteries
▪ The sound graph records the closing of
o This time period, when blood moves valves caused by blood flow
from the ventricles into the arteries
4. Period of isovolumetric relaxation
o STEP 4
o Ventricular repolarization, represented
by the T wave of an ECG, leads to
ventricular diastole
 ▪ As heart rate increases during
exercise, atrial contraction is
important for ventricular filling
because less time is available for
passive ventricular filling
▪ Therefore, it is during exercise that
the pumping action of the atria
becomes important for maintaining
the pumping efficiency of the heart
- Ventricular Systole: Period of
Isovolumetric Contract
o During the previous ventricular diastole,
the ventricles were filled with 120–130
mL of blood
o The volume of blood in the ventricles at
this point is the end-diastolic volume
(EDV)
o As the ventricles begin to contract,
ventricular pressure rapidly increases,
resulting in closure of the AV valves
▪ Ventricular volume does not change
during the period of isovolumetric
contraction because all the heart
valves are closed
- Ventricular Systole: Period of Ejection
o As soon as ventricular pressures exceed
the pressures in the aorta and pulmonary
Events Occurring During the Cardiac Cycle trunk, the semilunar valves open
The cardiac cycle is divided into five time periods (top). This graph ▪ The aortic semilunar valve opens at
represents several events that occur during the cardiac cycle. Each panel
represents a different aspect of cardiac function. From top to bottom:
approximately 80 mm Hg ventricular
Panel 1 represents the electrocardiogram; panel 2 represents pressure pressure, whereas the pulmonary
changes for the left atrium (blue line), left ventricle (black line), and semilunar valve opens at
aorta (red line); panel 3 represents the left ventricular volume curve;
and panel 4 represents heart sounds.
approximately 8 mm Hg
• Although the pressures are
different, both valves open at
- Atrial Systole and Active Ventricular nearly the same time
Filling o As blood flows from the ventricles during
o Before the cardiac cycle begins, all the period of ejection, the left ventricular
chambers are relaxed and blood is pressure continues to climb to
flowing from the veins into the atria and approximately 120 mm Hg, and the right
passively into the ventricles ventricular pressure increases to
▪ Most of ventricular filling occurs approximately 25 mm Hg
during this time ▪ The larger left ventricular pressure
▪ When the atria contract, active causes blood to flow throughout the
ventricular filling occurs as the force body (systemic circulation), whereas
of atrial contraction “tops off” the the lower right ventricular pressure
ventricles causes blood to flow through the
o Under most conditions, the atria function lungs (pulmonary circulation)
primarily as reservoirs, and the ventricles ▪ It is important to note that even
can pump sufficient blood to maintain though the pressure generated by the
homeostasis even if the atria do not left ventricle is much higher than that
contract at all of the right ventricle, the amount of
▪ During exercise, however, the heart blood pumped by each is almost the
pumps 300–400% more blood than same
during rest
 o During the first part of ejection, blood Heart Sounds
flows rapidly out of the ventricles - The pumping heart produces distinct sounds,
▪ Toward the end of ejection, as revealed by using a stethoscope
ventricular pressure decreases due to o These sounds are best heard by applying
reduced blood flow, despite the stethoscope at particular sites in
continued ventricular contraction relation to the heart valves
▪ By the end of ejection, the volume of - First heart sound
blood in the ventricles has decreased o A low-pitched sound, often described as
to 50–60 mL “lubb”
▪ The volume of blood remaining in o It occurs at the beginning of ventricular
the ventricles at the end of systole and is caused by vibration of the
ventricular systole is called the end- atrioventricular valves and surrounding
systolic volume (ESV) fluid as the valves close
- Ventricular Diastole: Period of - Second heart sound
Isovolumetric Relaxation o A higher-pitched sound often described
o Completion of the T wave results in as “dupp”
ventricular repolarization and relaxation o It occurs at the beginning of ventricular
o The already decreasing ventricular diastole and results from closure of the
pressure falls very rapidly as the aortic and pulmonary semilunar valves
ventricles suddenly relax as ventricular - Systole is therefore approximately the time
diastole begins between the first and second heart sounds
o When the ventricular pressures fall below - Diastole, which lasts somewhat longer, is
the pressures in the aorta and pulmonary approximately the time between the second
trunk, the recoil of the elastic arterial heart so und and the next first heart sound
walls, which were stretched during the
period of ejection, forces the blood to
flow back toward the ventricles, thereby
closing the semilunar valves
o Ventricular volume does not change
during the period of isovolumetric
relaxation because all the heart valves are
closed at this time
- Ventricular Diastole: Passive Ventricular
Filling
o The relaxed atria were filling with blood
during ventricular systole and the period
of isovolumetric relaxation
o During ventricular diastole, as ventricular
pressure drops below atrial pressure, the
atrioventricular valves open and allow Location of the Heart Valves in the Thorax
Surface markings of the heart in the male. The positions of
blood to flow from the filled atria into the the four heart valves are indicated by blue ellipses, and the
ventricles sites where the sounds of the valves are best heard with the
▪ At this point the atria and ventricles stethoscope are indicated by pink circles.
are relaxed - Faint third heart sound
o Blood flows from the area of higher o Can be heard in some normal people,
pressure in the veins and atria toward the particularly those who are thin and young
area of lower pressure in the relaxed o It is caused by blood flowing in a
ventricles turbulent fashion into the ventricles, and
o Most ventricular filling occurs during the it can be detected near the end of the first
first one-third of ventricular diastole one-third of diastole, during passive
o At the end of passive ventricular filling, ventricular filling
the ventricles are approximately 70%
filled
Aortic Pressure Curve
- The elastic walls of the aorta are stretched as
blood is ejected into the aorta from the left
ventricle
- Aortic pressure remains slightly below
ventricular pressure during this period of
ejection
o As blood leaves the ventricles, the
pressure in the ventricles begins to
decrease, even as the ventricles continue
to contract
▪ Similarly, pressure within the aorta
decreases as well
o As ventricular pressure drops below that
in the aorta, blood flows back toward the
ventricle because of the elastic recoil of
the aorta
▪ As the blood flows back toward the Aortic Pressure Curve
left ventricle, the aortic semilunar
valve closes
o Pressure within the aorta rises slightly at
this point
▪ This sudden change in aortic
pressure results in
a dicrotic notch, or incisura in the
aortic pressure curve
• The term dicrotic means
“double-beating”
• When increased pressure caused
by recoil is large, a double pulse
can be felt
o Aortic pressure then gradually falls
throughout the rest of ventricular diastole
as blood flows through the peripheral Cardiac Cycle: Pressure in the Heart
vessels
▪ When aortic pressure has fallen to
approximately 80 mm Hg, the
ventricles again contract, forcing
blood once more into the aorta
- Many of us have had our blood pressure
measured during a medical exam
o Blood pressure measurements
performed for clinical purposes reflect
the pressure changes that occur in the
aorta rather than in the left ventricle
The blood pressure in the aorta
fluctuates between systolic pressure,
which is about 120 mm Hg, and
diastolic pressure, which is about 80
mm Hg, for the average young adult at
rest
 MEAN ARTERIAL Cardiac Output
- Equal to heart rate times stroke volume
BLOOD PRESSURE
CO = HR x SV
Blood pressure is necessary to move the
o Heart rate (HR)
blood and therefore is critical to the maintenance of
▪ The number of times the heart beats
homeostasis. Blood flows from areas of higher
(contracts) per minute
pressure to areas of lower pressure. For example,
o Stroke volume (SV)
during one cardiac cycle, blood flows from the
▪ The volume of blood pumped during
higher pressure in the aorta, resulting from
each heartbeat (cardiac cycle)
contraction of the left ventricle, through the
o Stroke volume is equal to end-diastolic
systemic circulation, toward the lower pressure in
volume minus end-systolic volume
the relaxed right atrium.
▪ During diastole, blood flows from
the atria into the ventricles, and end-
Mean Arterial Pressure (MAP)
diastolic volume normally increases
- Slightly less than the average of the systolic
to approximately 125 mL
and diastolic pressure in the aorta
▪ After the ventricles partially empty
- It is proportional to cardiac output times
during systole, end-systolic volume
peripheral resistances
decreases to approximately 55 mL
- Cardiac Output (CO)
o Because stroke volume is equal to end-
o Minute volume
diastolic volume minus end-systolic
o The amount of blood pumped by the
volume, we can predict that stroke
heart per minute
volume is equal to 70 mL (125 − 55)
- Peripheral Resistance (PR)
- Stroke volume can be increased by increasing
o The total resistance aginst whih blood
end-diastolic volume or by decreasing end-
must be pumped
systolic volume
o During exercise, end-diastolic volume
MAP = CO x PR increases because of an increase
in venous return, which is the amount of
- Because mean arterial pressure is determined
blood returning to the heart from the
by both cardiac output and peripheral
systemic circulation
resustnce, changes to either can alter mean
o End-systolic volume decreases because
arterial pressure
the heart contracts more forcefully
▪ For example, stroke volume can
increase from a resting value of 70
mL to an exercising value of 115 mL
by increasing end-diastolic volume
to 145 mL and decreasing end-
systolic volume to 30 mL
- Cardiac output is also influenced by heart rate
(CO = HR × SV)
o Under resting conditions, the heart rate is
approximately 72 bpm, and the stroke
volume is approximately 70 mL/beat,
although these values can vary
considerably from person to person
o Therefore, the cardiac output is
Factors Affecting Mean Arterial Pressure
Mean arterial pressure is regulated by controlling cardiac output and
peripheral resistance.

- Changes in heart rate and stroke volume will

result in changes in cardiac output
 o For example, during exercise, the heart
rate can increase to 190 bpm, and the
stroke volume can increase to 115 mL
o Consequently, cardiac output is

Cardiac Reserve
- The difference between cardiac output when a
person is at rest and maximum cardiac output
- The greater a person’s cardiac reserve, the
greater his or her capacity for doing exercise
o Exercise can greatly increase cardiac
reserve by increasing cardiac output
▪ In well-trained athletes, stroke
volume during exercise can increase
to over 200 mL/beat, resulting in
cardiac outputs of 40 L/min or more
- Cardiovascular disease and lack of exercise
can reduce cardiac reserve and affect a
person’s quality of life
 REGULATION OF THE o This relationship between preload and
stroke volume and it describes the
HEART relationship between changes in the
pumping effectiveness of the heart and
To maintain homeostasis, the amount of changes in preload
blood pumped by the heart must vary dramatically, ▪ Preload, or ventricular stretching, is
depending on the level of activity and the O2and directly related to venous return
nutrient needs of the body tissues. For example, (remember our water balloon analogy)
during exercise, cardiac output can increase several ▪ Venous return can decrease to a value
times over resting values to meet the needs of the as low as 2 L/min or increase to as
active tissues. Intrinsic and extrinsic regulatory much as 24 L/min
mechanisms control cardiac output. • Such drastic changes in venous
return have major effects on the
Intrinsic regulation results from the preload
heart’s normal functional characteristics and does - Afterload
not depend on either neural or hormonal regulation. o The pressure the contracting left ventricle
It functions whether the heart is in place in the body must produce to overcome the pressure in
or is removed and maintained outside the body the aorta and move blood into the aorta
under proper conditions. o s the ventricles contract, pressure
increases, eventually forcing open the
Extrinsic regulation involves neural and semilunar valves
hormonal control. Neural regulation of the heart ▪ Although the heart’s pumping
results from sympathetic and parasympathetic effectiveness is greatly influenced by
reflexes, and the major hormonal regulation comes relatively small changes in the
from epinephrine and norepinephrine secreted by preload, it is very insensitive to large
the adrenal medulla. changes in afterload
• Aortic blood pressure must
Intrinsic Regulation increase to more than 170 mm
- The force of contraction produced by cardiac Hg before it hampers the
muscle is related to the degree of stretch of ventricles’ ability to pump blood
the cardiac muscle cells - During exercise, skeletal muscle activity
o As venous return increases, end-diastolic greatly influences heart activity by altering
volume increases venous return and preload
o A greater end-diastolic volume increases o During exercise, blood vessels in
the stretch of the ventricular walls exercising skeletal muscles dilate and
o Preload allow more blood to flow through the
▪ The extent to which the ventricular vessels
walls are stretched ▪ The increased blood flow increases
▪ An increased preload increases O2 and nutrient delivery to the
cardiac output, and a decreased exercising muscles
preload decreases cardiac output o In addition, skeletal muscle contractions
- Starling law of the heart repeatedly compress veins and cause
o The length-versus-tension relationship in blood to flow more rapidly from the
cardiac muscle is similar to that in skeletal skeletal muscles toward the heart
muscle ▪ As blood flows rapidly through
▪ Skeletal muscle, however, is normally skeletal muscles and back to the
stretched to nearly its optimal length heart, venous return to the heart
before contraction, whereas cardiac increases, increasing the preload
muscle cells are not stretched to the ▪ The increased preload causes an
point at which they contract with a increased force of cardiac muscle
maximal force contraction, which increases stroke
o An increased preload causes the cardiac volume
muscle cells to contract with a greater ▪ The increase in stroke volume results
force and produce a greater stroke volume in increased cardiac output, and the
 volume of blood flowing to the ▪ Strong parasympathetic stimulation
exercising muscles increases can decrease the heart rate below
o When a person rests, venous return to the resting levels by at least 20–30 bpm,
heart decreases because arteries in the but it has little effect on stroke
skeletal muscles constrict and because volume
muscular contractions no longer • In fact, if venous return remains
repeatedly compress the veins constant while the heart is
▪ As a result, blood flow through inhibited by parasympathetic
skeletal muscles decreases, and stimulation, stroke volume can
preload and cardiac output decrease actually increase
▪ The longer time between heartbeats
Extrinsic Regulation allows the heart to fill to a greater
- The heart is innervated by capacity, resulting in an increased
both parasympathetic and sympathetic preload, which in turn increases
nerve fibers stroke volume
o They influence the pumping action of the o Acetylcholine, the neurotransmitter
heart by affecting both heart rate and produced by postganglionic
stroke volume parasympathetic neurons, binds to ligand-
o The influence of parasympathetic gated channels that cause plasma
stimulation on the heart is much less than membranes of cardiac muscle cells to
that of sympathetic stimulation become more permeable to K+
o Sympathetic stimulation can increase ▪ As a consequence, the membrane
cardiac output by 50–100% over resting hyperpolarizes
values, whereas parasympathetic ▪ Heart rate decreases because the
stimulation can cause only a 10–20% hyperpolarized membrane takes
decrease longer to depolarize to the point of
- Extrinsic regulation of the heart keeps blood an action potential
pressure, blood O2 levels, blood CO2 levels,
and blood pH within their normal ranges of
values
o For example, if blood pressure suddenly - Sympathetic Control
decreases, extrinsic mechanisms detect o Sympathetic innervation of the heart
the decrease and initiate responses that begins with preganglionic neurons that
increase cardiac output to bring blood originate in the thoracic region of the
pressure back into its normal range spinal cord
- Parasympathetic Control ▪ These neurons synapse with
o Parasympathetic nerve fibers that postganglionic neurons of the
innervate the heart are in the Vagus inferior cervical and upper thoracic
Nerves sympathetic chain ganglia, which
▪ Preganglionic fibers of the vagus project to the heart as cardiac
nerve extend from the brainstem to nerves
terminal ganglia within the wall of ▪ The postganglionic sympathetic
the heart, and postganglionic fibers neurons innervate the SA and AV
extend from the ganglia to the SA nodes, the coronary blood vessels,
node, AV node, coronary blood and the atrial and ventricular
vessels, and atrial myocardium myocardia
o Has an inhibitory influence on the heart, o Increases both the heart rate and the force
primarily by decreasing the heart rate of muscular contraction
▪ When a person is at rest, continuous ▪ In response to strong sympathetic
parasympathetic stimulation inhibits stimulation, the heart rate can
the heart to some degree increase to 250 or, occasionally,
▪ An increase in heart rate during 300 bpm
exercise results, in part, from ▪ Stronger contractions can also
decreased parasympathetic increase stroke volume
stimulation
 • The increased force of depolarization, so that the frequency of
contraction resulting from the action potentials increases
sympathetic stimulation causes a ▪ The effect of norepinephrine on the
lower end-systolic volume in the heart involves its association with
heart; therefore, the heart cell surface β-adrenergic receptors
empties to a greater extent • This combination causes a G
protein–mediated synthesis and
accumulation of cAMP in the
cytoplasm of cardiac muscle
cells
• Cyclic-AMP increases the
permeability of the plasma
membrane to Ca2+, primarily by
opening calcium channels in the
plasma membrane
o Increased sympathetic stimulation causes
coronary arteries to constrict to some
degree
▪ However, increased metabolism of
Baroreceptor and Chemoreceptor Reflexes cardiac muscle, in response to
Reflexes in response to changes in blood pressure, pH, blood O2, and blood sympathetic stimulation, allows
CO2levels help regulate the activity of the heart to maintain homeostasis.
Sensory neurons (green) carry action potentials from sensory receptors to the metabolic by-products to accumulate
medulla oblongata. Sympathetic (blue) and parasympathetic (red) neurons in cardiac muscle, which causes
exit the spinal cord or medulla oblongata and extend to the heart to regulate coronary blood vessels to dilate
its function. Epinephrine and norepinephrine (dotted green line) from the
adrenal gland also help regulate the heart’s action (SA = sinoatrial). ▪ The dilation effect of these
metabolites predominates
o The relationship between increased heart
rate and cardiac output is limited
▪ If the heart rate becomes too fast,
ventricular diastole does not last long - The difference between parasympathetic and
enough to allow complete ventricular sympathetic
filling, end-diastolic volume o The sympathetic nervous system (SNS)
decreases, and stroke volume releases the hormones (catecholamines -
actually decreases epinephrine and norepinephrine) to
▪ If the heart rate increases beyond a accelerate the heart rate
critical level, the strength of o The parasympathetic nervous system
contraction decreases, probably (PNS) releases the hormone acetylcholine
because metabolites accumulate in to slow the heart rate
cardiac muscle cells - Hormonal Control
▪ The heart’s ability to increase the o Epinephrine and norepinephrine released
cardiac output is limited to heart from the adrenal medulla can markedly
rates of 170–250 bpm in response to influence the heart’s pumping
intense sympathetic stimulation effectiveness
o Sympathetic stimulation of the ▪ Epinephrine has essentially the same
ventricular myocardium plays a effect on cardiac muscle as
significant role in regulating its norepinephrine, increasing the rate
contraction force when a person is at rest and force of heart contractions
▪ Maintains the strength of ventricular o The secretion of epinephrine and
contraction at a level approximately norepinephrine is controlled by
20% greater than it would be without sympathetic stimulation of the adrenal
sympathetic stimulation medulla; it occurs in response to
o Norepinephrine, the postganglionic increased physical activity, emotional
sympathetic neurotransmitter, increases excitement, or other stressful conditions
the rate and degree of cardiac muscle ▪ Many stimuli that increase
sympathetic stimulation of the heart
 also increase the release of
epinephrine and norepinephrine from
the adrenal medulla
▪ Epinephrine and norepinephrine
travel in the blood through the
vessels of the heart to the cardiac
muscle cells, where they bind to β-
adrenergic receptors and stimulate
cAMP synthesis
▪ Epinephrine takes a longer time to
act on the heart than sympathetic
stimulation does, but the effect lasts
longer
o Epinephrine and norepinephrine are very
similar neurotransmitters and hormones
▪ While epinephrine has slightly more
of an effect on your heart,
norepinephrine has more of an effect
on your blood vessels
▪ Both play a role in your body's
natural fight-or-flight response to
stress and have important medical
uses as well
 THE HEART AND Summary of the Baroreceptor Reflex
The baroreceptor reflex maintains homeostasis in response to changes in

HOMEOSTASIS blood pressure. (1) Blood pressure is within its normal range. (2) Blood
pressure increases outside the normal range, which causes homeostasis to be
disturbed. (3) Baroreceptors in the carotid arteries and aorta detect the
increase in blood pressure, and the cardioregulatory center in the brain alters
The pumping efficiency of the heart plays autonomic stimulation of the heart. (4) Heart rate and stroke volume decrease.
(5) These changes cause blood pressure to decrease. (6) Blood pressure
an important role in maintaining homeostasis. returns to its normal range; homeostasis is restored. Observe the responses to
Blood pressure in the systemic vessels must be high a decrease in blood pressure outside its normal range by following the red
arrows.
enough to allow nutrient and waste product
exchange across the walls of the capillaries and to - Changes in blood pressure stimulate
meet metabolic demands. In addition, the heart’s baroreceptors, which then communicate with
activity must be regulated because the metabolic control centers in the medulla oblongata
activities of the tissues change under such o Sensory neurons, which are primarily
conditions as exercise and rest. Reflexes help found in the glossopharyngeal (cranial
regulate the activity of the heart to maintain nerve IX) and vagus (cranial nerve X)
homeostasis. Baroreceptor reflexes regulate blood nerves, carry action potentials from the
pressure, and chemoreceptor reflexes help regulate baroreceptors to an area in the medulla
the heart’s activity. oblongata called the cardioregulatory
center, where sensory action potentials
Effect of Blood Pressure are integrated
- Baroreceptor Reflexes ▪ There are two parts to the
o Detect changes in blood pressure and cardioregulatory center:
lead to changes in heart rate and force of • The cardioacceleratory center
contraction increases heart rate
o Stretch receptors, the sensory receptors of • The cardioinhibitory center
the baroreceptor reflexes, are in the walls decreases heart rate
of certain large arteries, such as the o Action potentials then travel from the
internal carotid arteries and the aorta cardioregulatory center to the heart
o Baroreceptors measure blood pressure by through both the sympathetic and the
detecting the degree of stretch of blood parasympathetic divisions of the
vessel walls autonomic nervous system
- At normal blood pressures (80–120 mm Hg),
action potentials are sent from the
baroreceptors in the internal carotid arteries
and aorta to the medulla oblongata at a
relatively constant frequency
o When blood pressure rises, the arterial
walls are stretched farther, and the action
potential frequency at the baroreceptors
increases
o When blood pressure decreases, the
arterial walls are stretched to a lesser
extent, and the action potential frequency
decreases
o In response to elevated blood pressure,
the baroreceptor reflexes reduce
sympathetic stimulation and increase
parasympathetic stimulation of the heart,
causing the heart rate to slow
▪ Decreased blood pressure causes
decreased parasympathetic and
increased sympathetic stimulation of
the heart, resulting in an increased
heart rate and force of contraction
o Withdrawal of parasympathetic
stimulation is primarily responsible for
 increases in heart rate up to - The increased cardiac output causes greater
approximately 100 bpm blood flow through the lungs, where CO2 is
o Larger increases in heart rate, especially eliminated from the body
during exercise, result from sympathetic o This helps lower the blood CO2 level to
stimulation within its normal range, which increases
o The baroreceptor reflexes are blood pH
homeostatic because they keep the blood - Chemoreceptors primarily sensitive to blood
pressure within a narrow range of values O2 levels are found in the carotid and aortic
that is adequate to maintain blood flow to bodies
the tissues o These small structures are located near
large arteries close to the brain and heart,
Effect of pH, Carbon Dioxide, and Oxygen and they monitor blood flowing to the
- Chemoreceptor Reflexes brain and the rest of the body
o Help regulate the heart’s activity o A dramatic decrease in blood O2 levels,
o Chemoreceptors sensitive to changes in as occurs during asphyxiation, activates
blood pH and CO2levels are found in the the carotid and aortic body
medulla oblongata chemoreceptor reflexes
▪ A drop in blood pH, which is often o When all the regulatory mechanisms
due to a rise in CO2 decrease function together, large, prolonged
parasympathetic and increase decreases in blood O2 levels increase the
sympathetic stimulation of the heart, heart rate
resulting in increased heart rate and ▪ Low blood O2 levels also increase
force of contraction stimulation of respiratory
movements
▪ Increased inflation of the lungs
stimulates stretch receptors in the
lungs
• Action potentials from these
stretch receptors influence the
cardioregulatory center, which
causes the heart rate to increase
▪ The reduced O2 levels that exist at
high altitudes can cause an increase
in heart rate even when blood
CO2 levels remain low
• However, the carotid and aortic
body chemoreceptor reflexes are
more important in regulating
respiration and blood vessel
constriction than heart rate

Effect of Extracellular Ion Concentration

- The ions that affect cardiac muscle function
are the same ions that influence membrane
potentials in other electrically excitable
tissues, K+, Ca2+, and Na+
o However, cardiac muscle responds to
these ions differently than nerve or
Summary of the Chemoreceptor Reflex
The chemoreceptor reflex maintains homeostasis in response to changes in
skeletal muscle tissue does
blood CO2 and H+ concentrations (pH). (1) Blood pH is within its normal ▪ For example, the extracellular levels
range. (2) Blood pH increases outside the normal range. (3) Chemoreceptors
in the medulla oblongata detect increased blood pH. Control centers in the
of Na+ rarely deviate enough from
brain decrease sympathetic stimulation of the heart and adrenal medulla. (4) normal to significantly affect cardiac
Heart rate and stroke volume decrease, reducing blood flow to the lungs. (5)
These changes cause blood pH to decrease (as a result of increase in blood
muscle function
CO2). (6) Blood pH returns to its normal range; homeostasis is restored. - Excess extracellular K+ in cardiac tissue
Observe the responses to a decrease in blood pH outside its normal range by
following the red arrows.
causes the heart rate and stroke volume to
 decrease. A twofold increase in extracellular muscles before they decrease enough to
K+results in heart block, which is the loss of markedly influence the heart’s function
action potential conduction through the heart
o The excess K+ in the extracellular fluid Effect of Body Temperature
causes changes in the membrane - Under resting conditions, the temperature of
potential that lead to a decreased rate at cardiac muscle normally does not change
which action potentials are conducted dramatically, although alterations in
along cardiac muscle cells temperature influence the heart rate
▪ As the conduction rates decrease, - Small increases in cardiac muscle temperature
ectopic action potentials can occur cause the heart rate to speed up, and decreases
o Elevated blood levels of K+ can produce in temperature cause the heart rate to slow
enough ectopic action potentials to cause o example, during exercise or fever,
fibrillation increased heart rate and force of
▪ The membrane potential changes contraction accompany temperature
also results in less Ca2+ entering the elevations, but the heart rate drops under
sarcoplasm of the cell; thus, the conditions of hypothermia
strength of cardiac muscle - During heart surgery, body temperature is
contraction lessens sometimes reduced dramatically on purpose
▪ Overall, excess extracellular to slow the heart rate and other metabolic
K+ results in drastic loss of heart functions
function
- Although the extracellular concentration of
K+ is normally small, a reduction in
extracellular K+ causes the resting membrane
potential to become hyperpolarized; as a
consequence, it takes longer for the
membrane to depolarize to threshold
o Ultimately, the reduction in extracellular
K+ results in a decrease in heart rate
o The force of contraction is not affected,
however
- A rise in the extracellular concentration of
Ca2+ produces a greater force of cardiac
contraction because of a higher influx of
Ca2+ into the sarcoplasm during action
potential generation
o Elevated plasma Ca2+ levels have an
indirect effect on heart rate because they
reduce the frequency of action potentials
in nerve fibers, thus reducing sympathetic
and parasympathetic stimulation of the
heart
o Generally, elevated blood Ca2+ levels
lower the heart rate
- A low blood Ca2+ level increases the heart
rate, although the effect is imperceptible until
blood Ca2+ levels are reduced to
approximately one-tenth of their normal value
o The reduced extracellular Ca2+ levels
cause Na+ channels to open, which
allows Na+ to diffuse more readily into
the cell, resulting in depolarization and
action potential generation
o However, reduced Ca2+ levels usually
cause death due to tetany of skeletal
 EFFECTS OF AGING abnormally. The aortic semilunar valve is
especially likely to become stenosed, but other
ON THE HEART heart valves, such as the bicuspid valve, may
become either stenosed or incompetent.
Gradual changes in heart function normally The atrophy and replacement of cells of the
occur with aging. These age-related changes are left bundle branch and a decrease in the number of
minor under resting conditions but become more SA node cells alter the electrical conducting system
significant in response to exercise or other age- of the heart and lead to a higher rate of cardiac
related diseases. Under resting conditions, the arrhythmias in elderly people.
mechanisms that regulate the heart compensate
The enlarged and thickened cardiac muscle,
effectively for most of the age-related changes.
especially in the left ventricle, requires more O2 to
pump the same amount of blood pumped by a
Hypertrophy of the left ventricle is a younger heart. This change is not significant unless
common age-related change. This appears to result the coronary circulation is diminished by coronary
from a gradual increase in the pressure in the aorta, artery disease. However, the development of
against which the left ventricle must pump blood, coronary artery disease is age-related, as is
and a gradual increase in the stiffness of cardiac congestive heart disease. Approximately 10% of
muscle tissue. The elevated aortic pressure results elderly people over 80 have congestive heart
from a gradual reduction in arterial elasticity, failure, and a major contributing factor is coronary
leading to increased stiffness of the aorta and other artery disease. Because of age-related changes in
large arteries. Myocardial cells accumulate lipids, the heart, many elderly people are limited in their
and the number of collagen fibers increases in ability to respond to emergencies, infections, blood
cardiac tissue. These changes make the cardiac loss, and stress.
muscle tissue stiffer and less compliant. The
increased volume of the left ventricle can
Exercise has many beneficial effects on the
sometimes result in higher left atrial pressure and
heart. Regular aerobic exercise improves the
increased pulmonary capillary pressure. This can
heart’s functional capacity at all ages, provided the
cause pulmonary edema and a tendency for older
person has no other conditions that cause the extra
people to feel out of breath when they exercise
workload on the heart to be harmful.
strenuously.
The maximum heart rate gradually
declines, as can be roughly predicted by the
following formula:

The rate at which cardiac muscle breaks

down ATP increases, and the rate of Ca2+ transport
decreases. The maximum rate at which cardiac
muscle can carry out aerobic respiration also
decreases. In addition, the degree to which
epinephrine and norepinephrine can increase the
heart rate declines. These changes lead to longer
contraction and relaxation times for cardiac muscle
and a decrease in the maximum heart rate. Both the
resting and maximum cardiac outputs slowly
decline as people age; by 85 years of age, the
cardiac output may have decreased by 30–60%.
Age-related changes also occur in the
connective tissue of the heart valves. The
connective tissue becomes less flexible, and
Ca2+ deposits increase. The result is an increased
tendency for the heart valves to function
 2. The inner surfaces of the atria are mainly
SUMMARY smooth. The auricles have muscular ridges
called pectinate muscles.
3. The ventricles have ridges called
20.1 Functions of the Heart trabeculae carneae.
The heart produces the force that causes blood to
circulate. External Anatomy and Coronary Circulation
1. Each atrium has a flap called an auricle.
2. The coronary sulcus separates the atria from
20.2Size, Shape, and Location of the Heart the ventricles. The interventricular grooves
1. The heart is approximately the size of a separate the right and left ventricles.
closed fist and is shaped like a blunt 3. The inferior and superior venae cavae and
cone. the coronary sinus enter the right atrium.
2. The heart lies obliquely in the The four pulmonary veins enter the left
mediastinum, with its base directed atrium.
posteriorly and slightly superiorly and its 4. The pulmonary trunk exits the right
apex directed anteriorly, inferiorly, and to ventricle, and the aorta exits the left
the left. ventricle.
3. The base is deep to the second intercostal 5. Coronary arteries branch off the aorta to
space, and the apex extends to the fifth supply the heart. Blood returns from the
intercostal space. heart tissues to the right atrium through the
coronary sinus and cardiac veins.
20.3 Anatomy of the Heart
The heart consists of two atria and two ventricles.
Heart Chambers and Valves
Pericardium 1. The interatrial septum separates the atria
1. The pericardium is a sac that surrounds the from each other, and the interventricular
heart and consists of the fibrous pericardium septum separates the ventricles.
and the serous pericardium. 2. The tricuspid valve separates the right
2. The fibrous pericardium helps hold the heart atrium and ventricle. The bicuspid valve
in place. separates the left atrium and ventricle. The
3. The serous pericardium reduces friction as chordae tendineae attach the papillary
the heart beats. It consists of the following muscles to the atrioventricular valves.
parts: 3. The semilunar valves separate the aorta and
▪ The parietal pericardium lines the pulmonary trunk from the ventricles.
fibrous pericardium.
▪ The visceral pericardium lines the 20.4 Route of Blood Flow Through the Heart
exterior surface of the heart. 1. Blood from the body flows through the
▪ The pericardial cavity lies between the right atrium into the right ventricle and
parietal and visceral pericardia and is then to the lungs.
filled with pericardial fluid, which 2. Blood returns from the lungs to the left
reduces friction as the heart beats. atrium, enters the left ventricle, and is
pumped back to the body.
Heart Wall
1. The heart wall has three layers: 20.5 Histology
▪ The outer epicardium (visceral
pericardium) provides protection Heart Skeleton
against the friction of rubbing organs. The fibrous heart skeleton supports the openings
▪ The middle myocardium is of the heart, electrically insulates the atria from
responsible for contraction. the ventricles, and provides a point of attachment
▪ The inner endocardium reduces the for heart muscle.
friction resulting from blood passing
through the heart. Cardiac Muscle
1. Cardiac muscle cells are branched and have
a centrally located nucleus. Actin and
 myosin are organized to form sarcomeres. 5. The plateau exists because voltage-gated
The sarcoplasmic reticulum and T tubules Ca2+ channels remain open.
are not as organized as in skeletal muscle. 6. The rapid phase of repolarization results
2. Cardiac muscle cells are joined by from closure of the voltage-gated
intercalated disks, which allow action Ca+ channels and the opening of many
potentials to move from one cell to the next. voltage-gated K+ channels.
Thus, cardiac muscle cells function as a unit. 7. The entry of Ca2+ into cardiac muscle cells
3. Cardiac muscle cells have a slow onset of causes Ca2+ to be released from the
contraction and a prolonged contraction time sarcoplasmic reticulum to trigger
caused by the length of time required for contractions.
Ca2+ to move to and from the myofibrils.
4. Cardiac muscle is well supplied with blood Autorhythmicity of Cardiac Muscle
vessels that support aerobic respiration. 1. Cardiac pacemaker muscle cells are
5. Cardiac muscle aerobically uses glucose, autorhythmic because of the spontaneous
fatty acids, and lactate to produce ATP for development of a pacemaker potential.
energy. Cardiac muscle does not develop a 2. The pacemaker potential results from the
significant oxygen deficit. movement of Na+ and Ca2+ into the
pacemaker cells.
Conducting System 3. Ectopic foci are areas of the heart that
1. The SA node and the AV node are in the regulate heart rate under abnormal
right atrium. conditions.
2. The AV node is connected to the bundle
branches in the interventricular septum by Refractory Periods of Cardiac Muscle
the AV bundle. Cardiac muscle has a prolonged depolarization
3. The bundle branches give rise to Purkinje and thus a prolonged refractory period, which
fibers, which supply the ventricles. allows time for the cardiac muscle to relax before
4. The SA node is made up of small-diameter the next action potential causes a contraction.
cardiac muscle cells that initiate action
potentials, which spread across the atria and Electrocardiogram
cause them to contract. 1. An ECG records only the electrical activities
5. Action potentials are slowed in the AV of the heart.
node, allowing the atria to contract and ▪ Depolarization of the atria produces
blood to move into the ventricles. Then the the P wave.
action potentials travel through the AV ▪ Depolarization of the ventricles
bundles and bundle branches to the Purkinje produces the QRS complex.
fibers, causing the ventricles to contract, Repolarization of the atria occurs
starting at the apex. The AV node is also during the QRS complex.
made up of small-diameter cardiac muscle ▪ Repolarization of the ventricles
fibers. produces the T wave.
2. Based on the magnitude of the ECG waves
20.6 Electrical Properties and the time between waves, ECGs can be
used to diagnose heart abnormalities.
Action Potentials
1. After depolarization and partial 20.7 Cardiac Cycle
repolarization, a plateau is reached, during 1. The cardiac cycle involves repetitive
which the membrane potential only slowly contraction and relaxation of the heart
repolarizes. chambers.
2. The movement of Na+ through the voltage- 2. Blood moves through the circulatory
gated Na+ channels causes depolarization. system from areas of higher pressure to
3. During depolarization, voltage-gated areas of lower pressure. Contraction of
K+ channels close, and voltage-gated the heart produces the pressure.
Ca2+ channels begin to open. 3. The cardiac cycle is divided into five
4. Early repolarization results from closure of periods:
the voltage-gated Na+channels and the
opening of some voltage-gated K+ channels.
 ▪ Active ventricular filling results Aortic Pressure Curve
when the atria contract and pump 1. Contraction of the ventricles forces blood
blood into the ventricles. into the aorta, producing the peak systolic
▪ Although the ventricles are pressure.
contracting, during the period of 2. Blood pressure in the aorta falls to the
isovolumetric contraction, diastolic level as blood flows out of the
ventricular volume does not aorta.
change because all the heart 3. Elastic recoil of the aorta maintains pressure
valves are closed. in the aorta and produces the dicrotic notch
▪ During the period of ejection, the and dicrotic wave.
semilunar valves open, and blood
is ejected from the heart. 20.8 Mean Arterial Blood Pressure
▪ Although the heart is relaxing, 1. Mean arterial pressure is the average
during the period of isovolumetric blood pressure in the aorta. Adequate
relaxation, ventricular volume blood pressure is necessary to ensure
does not change because all the delivery of blood to the tissues.
heart valves are closed. 2. Mean arterial pressure is proportional to
▪ Passive ventricular filling results cardiac output (amount of blood
when blood flows from the higher pumped by the heart per minute) times
pressure in the veins and atria to peripheral resistance (total resistance to
the lower pressure in the relaxed blood flow through blood vessels).
ventricles. 3. Cardiac output is equal to stroke
volume times heart rate.
Events Occurring During the Cardiac Cycle 4. Stroke volume, the amount of blood
1. Most ventricular filling occurs when blood pumped by the heart per beat, is equal
flows from the higher pressure in the veins to end-diastolic volume minus end-
and atria to the lower pressure in the relaxed systolic volume.
ventricles. ▪ Venous return is the amount
2. Contraction of the atria completes of blood returning to the
ventricular filling. heart. Increased venous
3. Contraction of the ventricles closes the AV return increases stroke
valves, opens the semilunar valves, and volume by increasing end-
ejects blood from the heart. diastolic volume.
4. The volume of blood in a ventricle just ▪ Increased force of
before it contracts is the end-diastolic contraction increases stroke
volume. The volume of blood after volume by decreasing end-
contraction is the end-systolic volume. systolic volume.
5. Relaxation of the ventricles results in the 5. Cardiac reserve is the difference
closing of the semilunar valves, the opening between resting and exercising cardiac
of the AV valves, and the movement of output.
blood into the ventricles.

Heart Sounds 20.9 Regulation of the Heart

1. Closure of the atrioventricular valves
produces the first heart sound. Intrinsic Regulation
2. Closure of the semilunar valves produces the 1. Venous return is the amount of blood that
second heart sound. returns to the heart during each cardiac cycle.
3. Turbulent flow of blood into the ventricles 2. The Starling law of the heart describes the
can be heard in some people, producing a relationship between preload and the stroke
third heart sound. volume of the heart. An increased preload
causes the cardiac muscle cells to contract
with a greater force and produce a greater
stroke volume.
Extrinsic Regulation 4. All regulatory mechanisms functioning
1. The cardioregulatory center in the medulla together in response to low blood pH, high
oblongata regulates parasympathetic and blood CO2, and low blood O2 levels usually
sympathetic nervous control of the heart. produce increased heart rate and
2. Parasympathetic stimulation is supplied by vasoconstriction. Decreased O2levels
the vagus nerve. stimulate an increase in heart rate indirectly
▪ Parasympathetic stimulation decreases by stimulating respiration, and the stretch of
heart rate. the lungs activates a reflex that increases
▪ Postganglionic neurons secrete sympathetic stimulation of the heart.
acetylcholine, which increases
membrane permeability to K+, Effect of Extracellular Ion Concentration
producing hyperpolarization of the 1. An increase or a decrease in extracellular
membrane. K+ decreases heart rate.
3. Sympathetic stimulation is supplied by the 2. Increased extracellular Ca2+ increases force
cardiac nerves. of contraction of the heart and decreases
▪ Sympathetic stimulation increases heart heart rate. Decreased Ca2+ levels produce
rate and force of contraction (stroke the opposite effect.
volume)
▪ Postganglionic neurons secrete Effect of Body Temperature
norepinephrine, which increases Heart rate increases when body temperature
membrane permeability to Na+ and increases, and it decreases when body temperature
Ca2+ and produces depolarization of the decreases.
membrane.
4. Epinephrine and norepinephrine are released
into the blood from the adrenal medulla as a 20.11 Effects of Aging on the Heart
result of sympathetic stimulation. 1. Aging results in gradual changes in
▪ The effects of epinephrine and heart function, which are minor under
norepinephrine on the heart are long- resting conditions but more significant
lasting, compared with those of neural during exercise.
stimulation. 2. Hypertrophy of the left ventricle is a
▪ Epinephrine and norepinephrine common age-related condition.
increase the rate and force of heart 3. The maximum heart rate declines so
contraction. that, by age 85, the cardiac output may
be decreased by 30–60%.
20.10 The Heart and Homeostasis 4. There is an increased tendency for
valves to function abnormally and for
Effect of Blood Pressure arrhythmias to occur.
1. Baroreceptors monitor blood pressure. 5. Because increased O2 consumption is
2. In response to a decrease in blood pressure, required to pump the same amount of
the baroreceptor reflexes increase blood, age-related coronary artery
sympathetic stimulation and decrease disease is more severe.
parasympathetic stimulation of the heart, 6. Exercise improves the functional
resulting in increased heart rate and force of capacity of the heart at all ages.
contraction.

Effect of pH, Carbon Dioxide, and Oxygen

1. Chemoreceptors monitor blood CO2, pH,
and O2 levels.
2. In response to increased CO2 and decreased
pH, medullary chemoreceptor reflexes
increase sympathetic stimulation and
decrease parasympathetic stimulation of the
heart.
3. Carotid body chemoreceptor receptors
stimulated by low O2 levels result in
decreased heart rate and vasoconstriction.