350px, Image showing the cardiac pacemaker or , the primary pacemaker within the electrical conduction system of the heart">SA node, the primary pacemaker within the electrical conduction system of the heart
The cardiac pacemaker is the heart's natural rhythm generator. It employs pacemaker Cell (biology), cells that produce electrical impulses, known as Cardiac action potential, cardiac action potentials, which control the rate of contraction of the
cardiac muscle, that is, the
heart rate
Heart rate is the frequency of the cardiac cycle, heartbeat measured by the number of contractions of the heart per minute (''beats per minute'', or bpm). The heart rate varies according to the body's Human body, physical needs, including the nee ...
. In most humans, these cells are concentrated in the
sinoatrial (SA) node, the primary pacemaker, which regulates the heart’s
sinus rhythm.
Sometimes a secondary pacemaker sets the pace, if the SA node is damaged or if the
electrical conduction system of the heart
The cardiac conduction system (CCS, also called the electrical conduction system of the heart) transmits the Cardiac action potential, signals generated by the sinoatrial node – the heart's Cardiac pacemaker, pacemaker, to cause the heart musc ...
has problems.
Cardiac arrhythmia
Arrhythmias, also known as cardiac arrhythmias, are irregularities in the heartbeat, including when it is too fast or too slow. Essentially, this is anything but normal sinus rhythm. A resting heart rate that is too fast – above 100 beat ...
s can cause
heart block, in which the contractions lose their rhythm. In humans, and sometimes in other animals, a mechanical device called an
artificial pacemaker
A pacemaker, also known as an artificial cardiac pacemaker, is an Implant (medicine), implanted medical device that generates Pulse (signal processing), electrical pulses delivered by electrodes to one or more of the Heart chamber, chambers of ...
(or simply "pacemaker") may be used after damage to the body's intrinsic conduction system to produce these impulses synthetically.
Control
Primary pacemaker
The
sinoatrial node
The sinoatrial node (also known as the sinuatrial node, SA node, sinus node or Keith–Flack node) is an ellipse, oval shaped region of special cardiac muscle in the upper back wall of the right atrium made up of Cell (biology), cells known as pa ...
(SA node) is the primary pacemaker of the heart. It is a region of cardiac muscle on the wall of the upper
right atrium
The atrium (; : atria) is one of the two upper chambers in the heart that receives blood from the circulatory system. The blood in the atria is pumped into the heart ventricles through the atrioventricular mitral and tricuspid heart valves.
...
near to the
superior vena cava
The superior vena cava (SVC) is the superior of the two venae cavae, the great venous trunks that return deoxygenated blood from the systemic circulation to the right atrium of the heart. It is a large-diameter (24 mm) short length vei ...
entrance. The cells that make up the SA node are specialized
cardiomyocytes known as pacemaker cells that can spontaneously generate
cardiac action potential
Unlike the action potential in skeletal muscle cells, the cardiac action potential is not initiated by nervous activity. Instead, it arises from a group of specialized cells known as pacemaker cells, that have automatic action potential generati ...
s. These signals are propagated through the heart's
electrical conduction system.
Only one percent of the heart muscle cells are conductive, the rest of the cardiomyocytes are
contractile.
The SA node controls the rate of contraction for the entire heart muscle because its cells have the quickest rate of spontaneous depolarization, thus they initiate action potentials the quickest. The action potential generated by the SA node passes down the
electrical conduction system of the heart
The cardiac conduction system (CCS, also called the electrical conduction system of the heart) transmits the Cardiac action potential, signals generated by the sinoatrial node – the heart's Cardiac pacemaker, pacemaker, to cause the heart musc ...
, and depolarizes the other potential pacemaker cells at the AV node to initiate action potentials before these other cells have had a chance to generate their own spontaneous action potential, thus they contract and propagate electrical impulses to the pace set by the cells of the SA node. This is the normal conduction of electrical activity in the heart.
The pacemaker cells are connected to neighboring contractile cells via
gap junction
Gap junctions are membrane channels between adjacent cells that allow the direct exchange of cytoplasmic substances, such small molecules, substrates, and metabolites.
Gap junctions were first described as ''close appositions'' alongside tight ...
s, which enable them to locally depolarize adjacent cells. Gap junctions allow the passage of positive cations from the depolarization of the pacemaker cell to adjacent contractile cells. This starts the depolarization and eventual action potential in contractile cells. Having cardiomyocytes connected via gap junctions allow all contractile cells of the heart to act in a coordinated fashion and contract as a unit. All the while being in sync with the pacemaker cells; this is the property that allows the pacemaker cells to control contraction in all other cardiomyocytes.
Cells in the SA node spontaneously
depolarize, ultimately resulting in contraction, approximately 100 times per minute. This native rate is constantly modified by the activity of
sympathetic and
parasympathetic
The parasympathetic nervous system (PSNS) is one of the three divisions of the autonomic nervous system, the others being the sympathetic nervous system and the enteric nervous system.
The autonomic nervous system is responsible for regulat ...
nerve fibers via the
autonomic nervous system
The autonomic nervous system (ANS), sometimes called the visceral nervous system and formerly the vegetative nervous system, is a division of the nervous system that operates viscera, internal organs, smooth muscle and glands. The autonomic nervo ...
, so that the average resting
heart rate
Heart rate is the frequency of the cardiac cycle, heartbeat measured by the number of contractions of the heart per minute (''beats per minute'', or bpm). The heart rate varies according to the body's Human body, physical needs, including the nee ...
in adult humans is about 70 beats per minute.
Secondary pacemaker
Impulses from the sinus node reach the
atrioventricular node
The atrioventricular node (AV node, or Aschoff-Tawara node) electrically connects the heart's atria and ventricles to coordinate beating in the top of the heart; it is part of the electrical conduction system of the heart. The AV node lies at the ...
which acts as the secondary pacemaker. The cells of the AV node normally discharge at about 40–60 beats per minute.
Atrioventricular junction
The atrioventricular node and the
Bundle of His
The bundle of His (BH) or His bundle (HB) ( "hiss"Medical Terminology for Health Professions, Spiral bound Version'. Cengage Learning; 2016. . pp. 129–.) is a collection of heart muscle cells specialized for electrical conduction. As part of ...
, a little further down, are located in the region separating the atria from the ventricles known as the atrioventricular junction. The Bundle of His transmits signals to the
bundle branches
The bundle branches, or Tawara branches, transmit cardiac action potentials (electrical signals) from the bundle of His to Purkinje fibers in heart ventricles. They are offshoots of the bundle of His and are important to the electrical conduc ...
, which send them on to the
Purkinje fibers
The Purkinje fibers, named for Jan Evangelista Purkyně, ( ; ; Purkinje tissue or subendocardial branches) are located in the inner ventricular walls of the heart, just beneath the endocardium in a space called the subendocardium. The Purki ...
. These will also produce a spontaneous cardiac action potential at a rate of 30–40 beats per minute, so if the SA and AV node both fail to function, these cells can also become pacemakers but with a much lower rate of conduction than either the primary or secondary pacemakers.
Generation of action potentials
There are three main stages in the generation of an action potential in a pacemaker cell. Since the stages are analogous to contraction of cardiac muscle cells, they have the same naming system. This can lead to some confusion as phases one and two are absent, leaving only phases zero, three, and four.
Phase 4Pacemaker potential
The key to the rhythmic firing of pacemaker cells is that, unlike
neuron
A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
s, these cardiomyocytes will slowly depolarize by themselves and do not need any outside innervation from the autonomic nervous system to fire action potentials.
In all other cells, the
resting potential
The relatively static membrane potential of quiescent cells is called the resting membrane potential (or resting voltage), as opposed to the specific dynamic electrochemical phenomena called action potential and graded membrane potential. The re ...
(-60mV to -70mV) is caused by a continuous outflow or "leak" of
potassium
Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...
ions through
ion channel
Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by Gating (electrophysiol ...
proteins
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, re ...
in the
membrane
A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. Bi ...
that surrounds the cells. However, in pacemaker cells, this potassium permeability (efflux) decreases as time goes on, causing a slow depolarization. In addition, there is a slow, continuous inward flow of
sodium
Sodium is a chemical element; it has Symbol (chemistry), symbol Na (from Neo-Latin ) and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 element, group 1 of the peri ...
, called the funny current, or
pacemaker current. These two relative ion concentration changes slowly depolarize (make more positive) the inside membrane potential (voltage) of the cell, giving these cells their pacemaker potential. When the membrane potential gets depolarized to about -40mV it has reached threshold (cells enter phase 0), allowing an action potential to be generated.
Phase 0Upstroke
Though much faster than the depolarization of phase 4, the upstroke in a pacemaker cell is slow compared to that in an
axon
An axon (from Greek ἄξων ''áxōn'', axis) or nerve fiber (or nerve fibre: see American and British English spelling differences#-re, -er, spelling differences) is a long, slender cellular extensions, projection of a nerve cell, or neuron, ...
.
The SA and AV node do not have fast sodium channels like neurons, and the depolarization is mainly caused by a slow influx of calcium ions. (The funny current also increases). Calcium enters the cell via voltage-sensitive calcium channels that open when the threshold is reached. This calcium influx produces the rising phase of the action potential, which results in the reversal of membrane potential to a peak of about +10mV. It is important to note that intracellular calcium causes muscular contraction in contractile cells, and is the effector ion. In heart pacemaker cells, phase 0 depends on the activation of
L-type calcium channels instead of the activation of voltage-gated fast sodium channels, which are responsible for initiating action potentials in contractile (non-pacemaker) cells. For this reason, the pacemaker action potential rising phase slope is more gradual than that of the contractile cell (image 2).
Phase 3Repolarization
The reversal of membrane potential triggers the opening of potassium leak channels, resulting in the rapid loss of potassium ions from the inside of the cell, causing repolarization (V
m gets more negative). The calcium channels are also inactivated soon after they open. In addition, as sodium channels become inactivated, sodium permeability into the cell is decreased. These ion concentration changes slowly repolarize the cell to resting membrane potential (-60mV). Another important note at this phase is that ionic pumps restore ion concentrations to pre-action potential status. The
sodium-calcium exchanger ionic pump works to pump calcium out of the
intracellular space, thus effectively relaxing the cell. The
sodium/potassium pump restores ion concentrations of sodium and potassium ions by pumping sodium out of the cell and pumping (exchanging) potassium into the cell. Restoring these ion concentrations is vital because it enables the cell to reset itself and enables it to repeat the process of spontaneous depolarization leading to activation of an action potential.
Clinical significance
Damage to the SA node
If the SA node does not function, or the impulse generated in the
SA node is blocked before it travels down the electrical conduction system, a group of cells further down the heart will become its pacemaker.
This center is typically represented by cells inside the
atrioventricular node
The atrioventricular node (AV node, or Aschoff-Tawara node) electrically connects the heart's atria and ventricles to coordinate beating in the top of the heart; it is part of the electrical conduction system of the heart. The AV node lies at the ...
(AV node), which is an area between the
atria and
ventricles, within the
atrial septum. If the AV node also fails,
Purkinje fibers
The Purkinje fibers, named for Jan Evangelista Purkyně, ( ; ; Purkinje tissue or subendocardial branches) are located in the inner ventricular walls of the heart, just beneath the endocardium in a space called the subendocardium. The Purki ...
are occasionally capable of acting as the default or "escape" pacemaker.
Ectopic pacemaker

An
ectopic pacemaker also known as an ectopic focus or ectopic foci, is a group of
excitable cells that causes a premature heart beat outside the normally functioning SA node of the heart. It is thus a cardiac pacemaker that is ectopic, producing an ectopic beat. If chronic this can result in
arhythmias such as
tachycardia
Tachycardia, also called tachyarrhythmia, is a heart rate that exceeds the normal resting rate. In general, a resting heart rate over 100 beats per minute is accepted as tachycardia in adults. Heart rates above the resting rate may be normal ...
,
bradycardia
Bradycardia, also called bradyarrhythmia, is a resting heart rate under 60 beats per minute (BPM). While bradycardia can result from various pathological processes, it is commonly a physiological response to cardiovascular conditioning or due ...
, or
ventricular fibrillation
Ventricular fibrillation (V-fib or VF) is an abnormal heart rhythm in which the Ventricle (heart), ventricles of the heart Fibrillation, quiver. It is due to disorganized electrical conduction system of the heart, electrical activity. Ventricula ...
. An
artificial pacemaker
A pacemaker, also known as an artificial cardiac pacemaker, is an Implant (medicine), implanted medical device that generates Pulse (signal processing), electrical pulses delivered by electrodes to one or more of the Heart chamber, chambers of ...
may be used to counter this.
Artificial pacemakers
A
pacemaker is an artificial cardiac pacemaker, that is an
implanted medical device that generates electrical impulses delivered by electrodes to the chambers of the heart either the upper atria, or lower ventricles to cause the targeted chambers to contract and pump blood. By doing so, the artificial pacemaker takes over from the primary SA node pacemaker to regulate the function of the heart's electrical conduction system.
See also
*
Intrinsic cardiac nervous system
References
{{DEFAULTSORT:Cardiac Pacemaker
Cardiac anatomy
Cardiac electrophysiology