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''Das Reizleitungssystem des Säugetierherzens'' (English: "''The Conduction System of the Mammalian Heart''") is a scientific
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monogra ...
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 Doctorate of ...
. 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 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) catheter reco ...
". The monograph revealed the existence of the
atrioventricular node The atrioventricular node or AV 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 lower back section of t ...
and the function of
Purkinje cells Purkinje cells, or Purkinje neurons, are a class of GABAergic inhibitory neurons located in the cerebellum. They are named after their discoverer, Czech anatomist Jan Evangelista Purkyně, who characterized the cells in 1839. Structure T ...
. 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 R ...
and
Martin Flack Martin William Flack (20 March 1882 – 16 August 1931) was a British physiologist who co-discovered the sinoatrial node with Sir Arthur Keith in 1907. Flack later became demonstrator of physiology at the London Hospital and later a lecturer. H ...
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 t ...
, which subsequently led to their discovery of the
sinoatrial node The sinoatrial node (also known as the sinuatrial node, SA node or sinus node) is an oval shaped region of special cardiac muscle in the upper back wall of the right atrium made up of cells known as pacemaker cells. The sinus node is approxima ...
. Throughout the beginning of the 20th century, Tawara's monograph influenced the work of many cardiologists and it was later cited by
Willem Einthoven Willem Einthoven (21 May 1860 – 29 September 1927) was a Dutch doctor and physiologist. He invented the first practical electrocardiograph (ECG or EKG) in 1895 and received the Nobel Prize in Physiology or Medicine in 1924 for it ("for the ...
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 t ...
was slow, because of the long interval between
atrial The atrium ( la, ātrium, , entry hall) is one of 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 valves. There are two at ...
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, also known as inflammatory cardiomyopathy, is an acquired cardiomyopathy due to inflammation of the heart muscle. Symptoms can include shortness of breath, chest pain, decreased ability to exercise, and an irregular heartbeat. Th ...
(which led to the discovery of
Aschoff bodies In medicine, Aschoff bodies are nodules found in the hearts of individuals with rheumatic fever. They result from inflammation in the heart muscle and are characteristic of rheumatic heart disease. These nodules were discovered independently by Lu ...
), 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 a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monogra ...
.


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 t ...
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, are a class of GABAergic inhibitory neurons located in the cerebellum. They are named after their discoverer, Czech anatomist Jan Evangelista Purkyně, who characterized the cells in 1839. Structure T ...
s act as a pathway for the "atrioventricular connecting system". # The atrioventricular connecting system starts in the
atrioventricular node The atrioventricular node or AV 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 lower back section of t ...
, moves into the
fibrocartilaginous Fibrocartilage consists of a mixture of white fibrous tissue and cartilaginous tissue in various proportions. It owes its inflexibility and toughness to the former of these constituents, and its elasticity to the latter. It is the only type of ...
portion of the
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interat ...
(Bundle of His), divides into defined left and right
bundle branch The bundle branches, or Tawara branches, are offshoots of the bundle of His in the heart's ventricle. They play an integral role in the electrical conduction system of the heart by transmitting cardiac action potentials from the bundle of His to ...
es, and descends into the terminal ends of the
Purkinje fibers The Purkinje fibers (; often incorrectly ; 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 Purkinje fibers are specia ...
. 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 R ...
, 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 t ...
. 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 t ...
(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 and one of the oldest of its kind. It is also the world's highest-impact academic journal. It was founded in England in 1823. The journal publishes original research articles ...
'' 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 t ...
and his increasing doubts about its existence, Keith (with his student
Martin Flack Martin William Flack (20 March 1882 – 16 August 1931) was a British physiologist who co-discovered the sinoatrial node with Sir Arthur Keith in 1907. Flack later became demonstrator of physiology at the London Hospital and later a lecturer. H ...
) 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 or sinus node) is an oval shaped region of special cardiac muscle in the upper back wall of the right atrium made up of cells known as pacemaker cells. The sinus node is approxima ...
(1907) Encouraged by their initial success and inspired by Tawara's discovery of the
atrioventricular node The atrioventricular node or AV 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 lower back section of t ...
, Keith and Flack extended their studies and eventually discovered the
sinoatrial node The sinoatrial node (also known as the sinuatrial node, SA node or sinus node) is an oval shaped region of special cardiac muscle in the upper back wall of the right atrium made up of cells known as pacemaker cells. The sinus node is approxima ...
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 Willem Einthoven (21 May 1860 – 29 September 1927) was a Dutch doctor and physiologist. He invented the first practical electrocardiograph (ECG or EKG) in 1895 and received the Nobel Prize in Physiology or Medicine in 1924 for it ("for the ...
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. It is an electrogram of the heart which is a graph of voltage versus time of the electrical activity of the hear ...
. ; 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 R ...
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 for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interat ...
, 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 t ...
’ 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 t ...
. 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 (; often incorrectly ; 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 Purkinje fibers are specia ...
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 signals generated by the sinoatrial node – the heart's pacemaker, to cause the heart muscle to contract, and pump blood through ...


References

{{DEFAULTSORT:Das Reizleitungssystem des Saugetierherzens Cardiology