Introduction

The heart carries out the vital function of pumping oxygenated blood around the body, for
which it has to contract and relax in a coordinated fashion. This contraction process is
preceded by electrical excitation, which under normal conditions is initiated by the SA node
as an action potential. An action potential is the rapid sequence of changes in the membrane
potential, resulting in an electrical impulse. This electrical impulse then travels down through
the heart's electrical conduction system to cause myocardial contraction followed by
relaxation in an orderly fashion. There are two main classifications of cells in the heart to be
considered: cardiomyocytes and pacemaker cells. Each of these individual cell types has a
distinct pattern of action potentials divided into several distinct phases. A shared
characteristic common to both cell types is the third phase, designated as repolarization.
Repolarization defines the resetting of the electrochemical gradients of the cell to prepare for
a new action potential. The action potential (AP) of the working myocardium lasts for several
hundreds of milliseconds, with the delayed repolarization securing a refractory state for new
excitations throughout the entire contraction phase. Delayed repolarization in the human
myocardium relies mainly on the vast diversity of cardiac potassium channels but also on a
particular redundancy in the heart known as the "repolarization reserve," in which one current
takes over if another one should fail.

Cellular Level
The adult mammalian heart is comprised of many cell types. These include cardiomyocytes,
fibroblasts, endothelial cells, and perivascular cells. Of these, the cardiomyocytes occupy a
significant volume of the heart. Functionally these cardiomyocytes can be in turn
differentiated into general cardiomyocytes cells and Pacemaker cells. Also, for understanding
transmural dispersion of repolarization, the cardiomyocytes are classified as epicardial cells
(those near the surface), M cell, and endocardial cells (near the ventricular cavity).
Pacemaker cells are highly specialized myocardial cells with an intrinsic ability to depolarize
rhythmically and initiate an action potential.] The pacemaker cells are located primarily in the
SA and atrioventricular (AV) nodes, with some cells also in the bundle of His and Purkinje
fibers. Pacemaker cells possess a characteristic known as automaticity and initiate action
potentials on their own. This action potential is conducted down the cardiac conduction
system as an electrical impulse and also between one cardiomyocyte to another through gap
junctions. This conduction helps the heart to contract in a synchronized fashion
Conduction system: The SA node is located superiorly in the right atrium near the opening
of the superior vena cava. From the SA node, the depolarization current spread through the
right atrium via gap junctions and also passes to the left atrium via Bachmann's bundle. From
the SA node, the impulse pass to the AV node through the internodal fibers. The AV node's
location is also in the right atrium but inferiorly at the interatrial septum. The atria and the
ventricles are isolated electrically, and the electrical impulses can only pass from the atria to
the ventricles via the AV node. AV node conduction is characterized by a conduction delay,
which ensures that ventricular contraction occurs after the atria empty its blood into the
ventricles. From the AV node, a depolarization wave passes through the bundle of His,
located in the interventricular septum. From here passing via the two bundle branches and
Purkinje fibers, the action potential reaches the ventricular cardiomyocytes.
Organ Systems Involved
In contrast to the cardiac system, action potentials of the nervous system propagate through
similar mechanisms and can elicit contractions of skeletal muscles. However, cardiac action
potentials, specifically those of the pacemaker cells, possess automaticity.

Function
Cardiac action potentials and their associated repolarizations are vital in stimulating and
maintaining the heart's regular contractions, which is essential in maintaining perfusion to the
vital organs of the body.

Mechanism
The cardiac cells can only propagate action potentials because of an electrochemical potential
gradient across cellular membranes. Ions, mainly sodium (Na+), potassium (K+), and calcium
(Ca2+), are present in different concentrations inside the cells vs. their surrounding
environments. Sodium and calcium concentrations are more extracellular, while potassium is
present at a higher concentration inside the cell. Voltage-sensitive ion channels are available
on cellular membranes to facilitate the movement of these ions. The tendency of ions to move
down their chemical gradient and the tendency for charges to balance out across membranes
contributes to a net electrochemical potential that varies with the status of ion channels. The
term used for these variations in status is a phase. Cycles of these phases initiate when the
cell membranes reach a threshold potential. This threshold potential is different for
cardiomyocytes and pacemaker cells. Cells can reach threshold potential through stimulus by
either adjacent cells, or, if they are pacemaker cells, possess automaticity.

The Conducting System of the Heart

The cardiac conduction system is a collection of nodes and specialised conduction cells that
initiate and co-ordinate contraction of the heart muscle. It consists of:

 Sinoatrial node
 Atrioventricular node
 Atrioventricular bundle (bundle of His)
 Purkinje fibres

Overview of Heart Conduction

The following sequence of electrical events occurs during one full contraction of the heart
muscle:
  An excitation signal (an action potential) is created by the sinoatrial (SA) node.
 The wave of excitation spreads across the atria, causing them to contract.
 Upon reaching the atrioventricular (AV) node, the signal is delayed.
 It is then conducted into the bundle of His, down the interventricular septum.
 The bundle of His and the Purkinje fibres spread the wave impulses along the
ventricles, causing them to contract.

Components of the Cardiac Conduction System

Sinoatrial Node

The sinoatrial (SA) node is a collection of specialised cells (pacemaker cells), and is located
in the upper wall of the right atrium, at the junction where the superior vena cava enters.

These pacemaker cells can spontaneously generate electrical impulses. The wave of
excitation created by the SA node spreads via gap junctions across both atria, resulting
in atrial contraction (atrial systole) – with blood moving from the atria into the ventricles.

The rate at which the SA node generates impulses is influenced by the autonomic nervous
system:

 Sympathetic nervous system – increases firing rate of the SA node, and thus
increases heart rate.
 Parasympathetic nervous system – decreases firing rate of the SA node, and thus
decreases heart rate.

Atrioventricular Node

After the electrical impulses spread across the atria, they converge at the atrioventricular
node – located within the atrioventricular septum, near the opening of the coronary sinus.

The AV node acts to delay the impulses by approximately 120ms, to ensure the atria have
enough time to fully eject blood into the ventricles before ventricular systole.

The wave of excitation then passes from the atrioventricular node into the atrioventricular
bundle.

Atrioventricular Bundle

The atrioventricular bundle (bundle of His) is a continuation of the specialised tissue of the
AV node, and serves to transmit the electrical impulse from the AV node to the Purkinje
fibres of the ventricles.

It descends down the membranous part of the interventricular septum, before dividing into
two main bundles:
  Right bundle branch – conducts the impulse to the Purkinje fibres of the right
ventricle
 Left bundle branch – conducts the impulse to the Purkinje fibres of the left ventricle.

Purkinje fibers
The Purkinje fibers are specialized conducting fibers composed of electrically excitable cells.
They are larger than cardiomyocytes with fewer myofibrils and many mitochondria. They
conduct cardiac action potentials more quickly and efficiently than any of the other cells in
the heart's electrical conduction system

Excitation-contraction (E-C) coupling

Excitation-contraction (E-C) coupling refers to the series of events that link the action
potential (excitation) of the muscle cell membrane (the sarcolemma) to muscular contraction.
Although E-C coupling in myocardium is similar in many ways to skeletal muscle and
smooth muscle, there are also critical differences.

Excitation begins in the SA node, which is referred to as the pacemaker of the heart. The
excitation spreads across the atria and reaches the AV node, which temporarily slows the
contraction speed to allow blood to move from the atria to the ventricles.

Excitation-contraction coupling (ECC) is a physiological process that links excitation of

muscles by the nervous system to their mechanical contraction

Once the muscle is excited by a neuron, calcium is released from the sarcoplasmic reticulum,
and the calcium binds to troponin, thus coupling the excitation of the muscle with the
contraction of the muscle itself. Calcium binds to troponin after being released from the
sarcoplasmic reticulum

Ca prolongs the period of cardiac muscle cell depolarization before repolarization begins.
Contraction of cardiac muscle occurs due to the binding of the myosin head to ATP, which
pulls actin filaments to the center of the sarcomere, the mechanical force of contraction.