Studies of the electrical activity of the ventricles and the origin of the QRS complex

Historical events in the development of cardiac electrophysiology are described briefly. Observations before 1900 showed that electrical changes accompanied activity of muscle and nerve. Other studies showed that electrical activity of the heart produced voltage changes on the human torso. In 1903 Einthoven developed the string galvanometer which made measurement of electrocardiographic potentials much easier, more accurate and more common. The bases of understanding of arrhythmias were established by Lewis in the early 1900's. Soon thereafter Wilson devised practical and theoretical approaches to the human electrocardiogram which led to many further developments. Events before 1950 established the existence and mechanism of electrical activity in excitable cells. Studies of the origin of QRS began in about 1950, with studies of depolarization of the canine ventricle. Studies of the human ventricle followed. In the 70's it appeared possible to solve the electrocardiographic forward problem, prediction of electrocardiographic potentials from a knowledge of intracardiac events. That solution appeared possible because of new approaches to the associated physical and computational problems. Attempts to solve the forward problem at that time assumed that the cardiac generator (the boundary between resting and depolarized cells) was a uniform double layer generator. (The strength of the generator is constant everywhere along the boundary). Meanwhile physiologists and anatomists had worked out the mechanism of communication between cardiac cells. The cells are longer than they are wide, and each cell can depolarize contiguous cells. The connections between cells are predominantly at the ends of the cell and the longitudinal depolarization of a cardiac mass travels three times as fast as transverse depolarization. The generator is not uniform but is strongest parallel to the long axes of the cells. Many or most of those working in the field did not recognize the importance of the connections between cardiac cells in not only the pathway of excitation, but also the potentials produced as the cells depolarized. A number of experiments indicated that the uniform double layer assumption led to both qualitative and quantitative errors in prediction of fields generated by depolarization of cardiac muscle. These are reviewed. There are now alternatives to the uniform model which recognize the non-uniformity of the cardiac generators, particularly the axial model. The forward problem is unsolved but it appears possible that these newer models will make a solution possible.