Das Reizleitungssystem Des Säugetierherzens
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''Das Reizleitungssystem des Säugetierherzens'' (English: "''The Conduction System of the Mammalian Heart''") is a scientific
monograph A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
published in 1906 by
Sunao Tawara was a Japanese pathologist known for the discovery of the atrioventricular node. Tawara was born in Ōita Prefecture and studied at the Medical School, Imperial University of Tokyo in Tokyo, graduating in 1901 and receiving his Medical Doctor, ...
. It has been recognized by cardiologists as a monumental discovery, and a milestone in
cardiac electrophysiology Cardiac electrophysiology is a branch of cardiology and Basic Science, basic science focusing on the electrical activities of the heart. The term is usually used in clinical context, to describe studies of such phenomena by invasive (intracardiac ...
". The monograph revealed the existence of 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 ...
and the function of Purkinje cells. It was used by
Arthur Keith Sir Arthur Keith FRS FRAI (5 February 1866 – 7 January 1955) was a British anatomist and anthropologist, and a proponent of scientific racism. He was a fellow and later the Hunterian Professor and conservator of the Hunterian Museum of the ...
and Martin Flack as a detailed guide in their attempts to verify the existence of 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 ...
, which subsequently led to their discovery of 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 ...
. Throughout the beginning of the 20th century, Tawara's monograph influenced the work of many cardiologists and it was later cited by Willem Einthoven in his anatomical interpretation of the electrocardiogram.


Background

Prior to Tawara's discoveries, it was assumed that electrical conduction through 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 ...
was slow, because of the long interval between
atrial 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. ...
and ventricular contractions. The Swiss cardiologist Wilhelm His, Jr. assumed that the heart bundle was connected directly to the base of the ventricle, and physiologists incorrectly taught that the base of the ventricle contracted first, followed by the apex. However, Tawara postulated that ventricular contraction occurs in the opposite manner, with the apex contracting earlier than the base. He also believed that the heart's electrical conduction was not slow but rapid. Working under the guidance of his mentor,
Ludwig Aschoff Karl Albert Ludwig Aschoff (10 January 1866 – 24 June 1942) was a German physician and pathologist. He is considered to be one of the most influential pathologists of the early 20th century and is regarded as the most important German patholo ...
, Tawara performed a histological examination of 150 hearts with
myocarditis Myocarditis is inflammation of the cardiac muscle. Myocarditis can progress to inflammatory cardiomyopathy when there is associated ventricular remodeling and cardiac dysfunction due to chronic inflammation. Symptoms can include shortness of bre ...
(which led to the discovery of Aschoff bodies), and he began examining the atrioventricular bundle before embarking on a comprehensive study of the anatomy and histology of the heart's conduction system. The implications of his work were immediately recognized by Aschoff, who arranged for it to be published in the form of a
monograph A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
.


Discoveries

Tawara's monograph, titled "''Das Reizleitungssystem des Säugetierherzens''" (English: "'' The Conduction System of the Mammalian Heart''") was published in 1906. The most important discoveries are listed below: # 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 ...
is divided into 2 bundle branches that are connected with a fanlike group of “subendocardially scattered characteristic muscular bundles”. #
Purkinje cell Purkinje cells or Purkinje neurons, named for Czech physiologist Jan Evangelista Purkyně who identified them in 1837, are a unique type of prominent, large neuron located in the Cerebellum, cerebellar Cortex (anatomy), cortex of the brain. Wi ...
s act as a pathway for the "atrioventricular connecting system". # The atrioventricular connecting system starts in 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 ...
, moves into the
fibrocartilaginous Fibrocartilage consists of a mixture of white fibrous tissue and Cartilage, cartilaginous tissue in various proportions. It owes its inflexibility and toughness to the former of these constituents, and its Elasticity (physics), elasticity to the ...
portion of the
septum In biology, a septum (Latin language, Latin for ''something that encloses''; septa) is a wall, dividing a Body cavity, cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Hum ...
(Bundle of His), divides into defined left and right
bundle branch The bundle branches, or Tawara branches, transmit cardiac action potentials (electrical signals) from the bundle of His to Purkinje fibers in Ventricle (heart), heart ventricles. They are offshoots of the bundle of His and are important to the Ca ...
es, and descends into the terminal ends of 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 ...
. Tawara commented that the system represents a transporting or conducting pathway, and "because the pathway is not a ductal, but a continuously related protoplasmic cord, conduction of excitation impulses surely must take place there."


Influences

On 26 September 1905, shortly before the monograph was due to be published,
Ludwig Aschoff Karl Albert Ludwig Aschoff (10 January 1866 – 24 June 1942) was a German physician and pathologist. He is considered to be one of the most influential pathologists of the early 20th century and is regarded as the most important German patholo ...
wrote an article about Tawara's work. It was subsequently read by the Scottish cardiologist James Mackenzie and forwarded to anatomist
Arthur Keith Sir Arthur Keith FRS FRAI (5 February 1866 – 7 January 1955) was a British anatomist and anthropologist, and a proponent of scientific racism. He was a fellow and later the Hunterian Professor and conservator of the Hunterian Museum of the ...
, who was attempting to confirm the existence of 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 ...
. Despite putting in his best efforts, he failed to locate the structure. On 15 January 1906, Keith wrote a letter to Mackenzie and acknowledged his skepticism about its existence: "''I have given up the search for His' bundle—having come to the conclusion that there is not and never was any such thing.''” ; Verification of 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 ...
(1906) In response to Arthur Keith's skepticism, Mackenzie forwarded Aschoff's article about Tawara's findings, which stimulated Keith's renewal of his studies on the cardiac conduction system. Despite having written a letter to ''
The Lancet ''The Lancet'' is a weekly peer-reviewed general medical journal, founded in England in 1823. It is one of the world's highest-impact academic journals and also one of the oldest medical journals still in publication. The journal publishes ...
'' about his failure to locate 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 ...
and his increasing doubts about its existence, Keith (with his student Martin Flack) later reported that they had succeeded in locating the structure by following the detailed descriptions and figures in Tawara's monograph. In a paper published in ''The Lancet'' on 11 August 1906, they acknowledged the monograph's high degree of accuracy: :"We take this opportunity of clearly stating that although some of our observations are new our work is in the main but a verification of the accurate and complete monograph published recently by Tawara, a Japanese working in the laboratory of Professor Aschoff of Marburg." ; Discovery of 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 ...
(1907) Encouraged by their initial success and inspired by Tawara's discovery of 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 ...
, Keith and Flack extended their studies and eventually discovered 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 ...
in 1907. They wrote that they were examining other regions of the heart for "peculiar musculature" similar to the one discovered by Tawara. ; Theoretical basis for the electrocardiogram (1908) In 1908, the Dutch physiologist Willem Einthoven referred to Tawara’s monograph as the anatomical basis for interpreting the electrocardiogram. In his monograph, Tawara theorized about the velocity of the excitatory process in the conduction system and the mode of ventricular contraction. Together with his anatomic findings and physiological assumptions, it contributed to the rapid popularization of
electrocardiography Electrocardiography is the process of producing an electrocardiogram (ECG or EKG), a recording of the heart's electrical activity through repeated cardiac cycles. It is an electrogram of the heart which is a graph of voltage versus time of t ...
. ; Other influences and reviews In 1909, the American pathologist Lydia DeWitt created the first 3D wax model of the conduction system, using Tawara’s description as a guide. In 1911, the British cardiologist Thomas Lewis reviewed the auriculo-ventricular connection system and described Tawara's discoveries in ''Das Reizleitungssystem des Saugetierherzens'' as the "main advance" in knowledge about the system: :"The main advance was made by Tawara, working under the direction of Aschoff. In his book, ''Das Reizleitungssystem des Saugetierherzens'', a complete account of the junctional tissues was given, and the anatomy of the whole system and the connections with the network of Purkinje were described in great detail and in many species of animals. These observations upon the anatomy have received complete confirmation by the more recent writings." In his autobiography published in 1950,
Arthur Keith Sir Arthur Keith FRS FRAI (5 February 1866 – 7 January 1955) was a British anatomist and anthropologist, and a proponent of scientific racism. He was a fellow and later the Hunterian Professor and conservator of the Hunterian Museum of the ...
explained how he had systematically searched for Tawara's system to verify its components: :"I was able in heart after heart to verify the existence of Tawara’s system. The auricles, I found, were joined to the ventricles by an elaborate system which, beginning in a root like structure in the auricular
septum In biology, a septum (Latin language, Latin for ''something that encloses''; septa) is a wall, dividing a Body cavity, cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Hum ...
, ended as an arborescence in the ventricles. 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 ...
’ was but a small segment of the Tawara system." Acknowledging the significance and implications of these discoveries, Keith commented: "With the discovery of the conducting system of Tawara, heart research entered a new epoch." Shortly before his death, Wilhelm His, Jr. published a personal account about the discovery of 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 ...
. He noted that it took ten years before anatomists began to pay attention to the bundle, starting with the studies of Retzer and Brauning in 1903, followed by the "important work of Tawara" in 1906, and the subsequent discovery of the sinus node which completed the system. He credited Tawara for connecting the bundle with 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 ...
and for declaring it the heart's conduction system.


See also

*
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 ...


References

{{DEFAULTSORT:Das Reizleitungssystem des Saugetierherzens Cardiology